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A TEXT-BOOK OF BOTANY
 MACMILLAN AND CO., LIMITED
LONDON BOMBAY CALCUTTA
MELBOURNE

THE MACMILLAN COMPANY

NEW YORK BOSTON CHICAGO
ATLANTA SAN FRANCISCO

THE MACMILLAN CO. OF CANADA, LTD.

TORONTO
 TEXT-BOOK OF BOTANY

BY

DR. EDUARD STRASBURGER DR. FRITZ NOLL

PROFESSOR IN THE UNIVERSITY PROFESSOR IN THE AGRICULTURAL ACADEMY
OF BONN OF POPPELSDORF AND IN THE UNIVERSITY OF BONN

DR. HEINRICH SCHENCK DR. GEORGE KARSTEN

PROFESSOR IN THE TECHNICAL PROFESSOR IN THE UNIVERSITY
ACADEMY OF DARMSTADT OF BONN

THIRD ENGLISH EDITION

REVISED WITH THE EIGHTH GERMAN EDITION BY

W. H. LANG, M.B., D.Sc.

LECTURER IN BOTANY, UNIVERSITY OF GLASGOW

WITH 779 ILLUSTRATIONS, IN PART COLOURED

MACMILLAN AND CO., LIMITED

ST. MARTIN'S STREET, LONDON
1908
First English Edition, 1898

Second English Edition, 1903

Third English Edition, 1008

J.
 PREFATORY NOTE
THE first edition of the English translation of this text-book was

the work of Dr. H. 0. Porter, Assistant Instructor of Botany,

University of Pennsylvania. The proofs of this edition were

revised by Professor Seward, M.A., F.K.S. The second English
edition was based upon Dr. Porter's translation, which was revised
with the fifth German edition. The present edition has been

similarly revised throughout with the eighth German edition.

Such extensive changes, including the substitution of a completely

new section on Phanerogamia, have however been made in the
latter that seemed advisable to give in outline the history of
it

the English translation instead of retaining Dr. Porter's name

on the title page.
The official plants mentioned under the Natural Orders are
those of the British Pharmacoposia instead of those official in

Germany, Switzerland, and Austria, which are given in the

original. In making this alteration I have consulted Materia
Medica and Therapeutics, by J. Mitchell Bruce, M.A., LL.D., M.D.
The fact that a considerable portion of the original has been
rewritten, has necessitated the preparation of a new and extended
index. I am indebted to my friend, Mr. F. Tidd Pratt, for
assistance in this.
WILLIAM H. LANG.
GLASGOW, February 1908.

/\ ^ - ^v ^
^Vo s^x<l
L ' B R A R y
^^.
 CONTENTS
PAGE
INTRODUCTION 1

PART I. GENERAL BOTANY

SECTION I.' MORPHOLOGY

I. EXTERNAL MORPHOLOGY

The shoot .........

The development of form in the plant

.........
Metamorphosis of the shoot .
kingdom

. .
.

.
.

.
.

.
.11
18
.23
Leaves

The root ..........

Metamorphosis of leaves

Metamorphosis of the root

Members of independent origin
.

.
.

.
.

.
.

.
.

.
.

.
30
.43
45
.48
.49

II. INTERNAL MORPHOLOGY

The cell

Protoplasm
...
.........
(Histology and Anatomy)
52
54
Cell wall

Cell sap .........

........
Inclusions of the protoplasm . . . . . .
63
.73
79

Cell fusions
Tissues ..........
Ontogeny of the

Primary tissues
.
cell

......... . . . . . .
80
.93
97
100
.127
Secondary tissues

......
. . . . . . .

.151
Regeneration

......
. . . . . . . .

Phylogeny of the internal structure 152

Ontogeny of the internal structure 154
Structural deviations 165
 via BOTANY

SECTION II. PHYSIOLOGY

....
PA01

......
Physical and vital properties. General conditions of life 172
The stability of the plant body

......
178
Nutrition
Respiration
Growth .......
The phenomena
......of movement . . . .
186
239
247
263
Reproduction 296

PART II. SPECIAL BOTANY

A ...... SECTION I. CRYPTOGAMS

/'(^THALLOPHYTA
Bacteria
.......
.......
.

.......
Cyanophyceae
. . . .
329
331
337
Flagellata
Myxomycetes
Peridineae .......
.......
339
340
342

.......
Conjugatae 344

......
Diatomeae . . 347

.......
Heterocoutae 351

Characeae
.......
Chlorophyceae

.......
353
363

.......
Phaeophyceae 365

'
.......
Rhodophyceae

.......
Phycomycetes
373
378

.......
Eumycetes 386
O
.......
Lichcnes 415
6BRYOPHYTA

........ 421
/ -
fl
Hepaticae 425
__ Musci 431

......
438
/^PTERIDOPHYTA . .
(
Filicinae ... 443

....
. . . . .

.......
Eusporangiatae 443

.......
Leptosporangiatae
Equisetinae
Lycopodinae
......
FOSSIL CRYPTOGAMS
445
455
458
465

SECTION II. PHANEROGAMIA

TRANSITION FROM THE CRYPTOGAMS TO THE PHANEROGAMS

.......
'in
TABLE OF CLASSES, ORDERS, AND FAMILIES OF PHANEROGAMS 478
GYMNOSPERMAE 480
Development of the sexual generation 480
Cycadinae . . . . . 484
 CONTENTS ix

.......
......... PACK

Coniferae
Gnetinae .........
Ginkgoinae

...
.........
. 488
489
498

.......
FOSSIL GYMNOSPEKMS . 500
(

.........
ANGIOSPEKMAE

........
The angiospermic

..........
Inflorescence
Sexual generation
flower
501
501
509
513
Fruit

Helobiae
...
.......
tffG MONOCOTYLAK . .

.522
519
521

Glumiflorae . . . . . . . . .524
Spadicittorae . . . . . . . . . 530
Enantioblastae 536

....
. . .

Liliiflorae .536

....
.

Scitamineae . 546
Gynandrae
Cff DICOTYLAE
Choripetalae
Piperinae
...
....
.
.548
.651
.552
. 552
Juglandiflorae
Saliciflorae

Querciflcrae
...
.....
. . .

.
554
555
556
Urticinae . . 563
Ceutrospermae . . . 566

Polycarpicae
Rhoeadinae
Insectivorae
... .

. .
.571
.

.
585
591
Saxifraginae 591
Rosinorae . 593
Leguminosae . . 599
Gruinales
Tricoccae ...
...
. .

.615
610

Sapindinae
Frangulinae
Columniferae
... .
618
621
623
Cistiflorae
Passiflorinae

Opuntinae
...
.

.
.
.
627
630

631
Thymelaeinae .

Myrtiflorae . . .631
Umbelliflorae . 635

Hysterophyta
I/ SYMPETALAE .
- 645
645
Pentacyclicae .

Ericinae . .
645
. 646
Diospyrinae .

Primulinae . .
647

Tetracyclicae .

Contortae .
649
x BOTANY

......
.........
Tubiflorae .
PAOI
656

........
Personatae

.........
Rubiinae .

........
Campanulinae
Aggregatae
FOSSIL ANGIOSPERMS
. . . . . . . .668
661

675
679
688

.......
INDKX OF LITERATURE. 691

INDEX ........
SYSTEMATIC INDEX OF OFFICIAL AND POISONOUS PLANTS . . . 717

721
 INTRODUCTION

IT is customary to place all

living beings in either the animal or
vegetable kingdoms, but in reality a sharp boundary line between
animals and plants first becomes possible when they exhibit a
complicated structure. In those of more simple organisation all
distinctions disappear, and it becomes difficult to define the exact
limits of Botany and Zoology. This, in fact, could scarcely be
otherwise, as all the processes of life, in both the animal and vegetable
kingdoms, are dependent on the same substance, protoplasm. With
more complicated organisation, the specific differences increase, and
the characteristics distinguishing animal from vegetable life become
more obvious. For the present, it must be confessed, the recognition
of an organism, as an animal or a plant, is dependent upon its corre-
spondence with an abstract idea of what a plant or animal should
be, based on certain points of agreement between the members of
each class. A satisfactory basis for the separation of all living

organisms into the categories of animals or plants can only be

obtained when it is shown that all organisms distinguished as
animals are in reality genetically connected, and that a similar
connection exists between all plants. The proof of this can only be
arrived at through the THEORY OF EVOLUTION.
From the study of the fossil remains and impressions of animals
and plants, it has been established that in former epochs forms of
life differing from those of the present age existed on the earth.
It is also generally assumed that all living animals and plants have
been derived by gradual modification from previously existing forms.
This leads to the further conclusion that those organisms possess-
ing closely similar structure, which are united as species in a
genus, are in reality related to one another. It is also probable
that the union of corresponding genera into one family and of families
into higher groups serves to give expression to a real relationship

existing between them.

The presumable origin of a living organism from others previously
existing has been distinguished by HAECKEL ( ) as its phylogenetic
x
2 BOTANY

development or PHYLOGENY. He termed the series of changes

passed through by a living being in attaining its mature condition,
its ontogenetic development or ONTOGENY. The supposition, that
the successive steps in the ontogenetic development of an organism
correspond to those of its phylogenetic development, and that the
ontogeny of an organism is accordingly a more or less compk-te
repetition of its phylogeny, was asserted by FRITZ MULLER ( ), who
2

based his conclusions on the results of comparative research.

The idea of the gradual evolution of higher organisms from lower
was familiar to the Greek philosophers, but a scientific basis was first
given to this hypothesis in the last century. Through the work of
CHARLES DARWIN 3
( )
in particular, who accumulated evidence for a
reconsideration of the whole problem of organic evolution, the belief
in the immutability of species has been overturned.
DARWIN is THEORY OF SELECTION. In drawing
the author of the
his conclusions,he proceeded from the variability of living organisms,
as shown by the fact that the offspring neither exactly resemble
their parents nor each other. Further, he called attention to the
constant over-production of offspring, the majority of which must
inevitably be destroyed. If this were not so, and all the embryos

produced by a single pair attained their full development, they

would alone, in a few generations, completely cover the whole
surface of the earth. On account of insufficient space for all, the
different claimants are engaged in an uninterrupted struggle, in which
the victory is gained by those that, for any reason, have an advantage.
" "
Through this struggle for existence a selective process goes on
among the characters appearing in individual variations, and those
which under the conditions of life are in any way advantageous tend
to be preserved. In this manner DARWIN arrived at the supposition
of a process of NATURAL SELECTION, which is the essential of his
theory. Newly developed peculiarities arising from individual vari-
ability must be inherited in order to become permanent characteristics
of a later generation. DARWIN sought in the experience of breeders
evidence that such characters are inherited. The breeder selects indi-
viduals presenting any desired characters for the purpose of breeding,
and has thus formed the races of domesticated animals and cultivated
plants. These have often departed so widely from their wild
ancestral forms that the latter are not certainly known. Just as
in artificial selection, natural selection, although unconsciously, ac-

complishes this result. As individual peculiarities may be developed

by careful breeding and rendered permanent, so by natural selection
those qualities which are advantageous in the struggle for existence
become more pronounced and are finally confirmed by heredity. By
the continued operation of natural selection, organisms must result,
which are, in the highest degree, fitted and adapted to their
environment. In this way the theory of selection seeks to explain as
 INTRODUCTION

due to natural causes that adaptability to the environment which is

such a striking characteristic of organic life. That the transitional

forms in this process of phylogenetic development no longer exist, is
accounted for in the theory of natural selection by the assumption
that the struggle for existence must necessarily have been most
severe between similar organisms. For similar organisms have
similar necessities,and the new and better-equipped forms must
ultimately prevail over the original less specialised organisms and
exterminate them.
Since the publication of Darwin's works many investigators have
laboured to advance and make clear our views on phylogeny.
Difficulties inapplying the results of artificial selection to the natural
process became evident, for one main condition of successful artificial
selection, the isolation of the organisms from which breeding is
taking place, is not fulfilled under natural conditions. Of late years
HUGO DE VRIES has endeavoured to obtain an insight into the laws
of phylogenetic development by systematic cultivation of particular
4
plants. It would appear from such cultures ( ) that the starting-
"
point for the origin of new species is not afforded by the fluctuating
variations," which continually occur, but by more marked variations
"
which have been termed " mutations ;
these mutations appeal-
suddenly and are strongly inherited. On the other hand it may be
said that a sharp line cannot be drawn between mutations and
fluctuating variations. DE VRIES tended to assume the existence of
a development of the organic world due to original innate capabilities
of the living substance, and not dependent on selection. The origin
of the large subdivisions of the animal and vegetable kingdoms, the
5
"archetypes," would be due to this sort of evolution ( ). The
organisms have been, and are still, continually influenced by the
environment, and by their reaction to external conditions have
become more or less directly adapted. In this way striking re-
semblances in external form have arisen between organisms living
under similar conditions although belonging to different archetypes ( 6 ).
Natural selection exercises a constant influence on the process and
tends to render species distinct by removing the less advantageous
variations.
If the higher organisms have been evolved from the lower, a

sharp distinction between plants and animais is excluded. For the

characters which are distinctive of animals and plants have appeared
in the course of the phylogenetic development of organisms, and
were at first wanting. The simplest organisms which now exist are
in all probability similar to those which formed the starting-point
of this development. The walls which surround the cells com-
posing the plant body, and the green chroma tophores within the
latter, have been cited as decisive indications of the vegetable
character of an organism. Surrounded by firm walls, the living
4 BOTANY

substance becomes more isolated, and, consequently, independence

of action in plants, as compared with animals, is diminished. By
means of the green colouring matter, plants have the power of
producing their own nutritive substances from certain constituents
of the air and water, and from the salts contained in the soil, and
are thus able to exist independently while animals are dependent,
;

and so for their very

directly or indirectly, for their nourishment,
existence, on plants. Almost all the other differences which dis-
tinguish plants from animals may be traced to the structure of
plants, or to the manner in which they obtain their food. Another
characteristic of plants the unlimited duration of their ontogenetic
is

development, which continuous, at the growing points, during

is

their whole life. That none of these criteria are alone sufficient
for distinguishing plants from animals is evident from the fact that
all the Fungi are devoid of green pigment, and, like animals, are

dependent on substances produced by green plants for their

nourishment. On the borderland of the two kingdoms, where all
other distinctions are wanting, phylogenetic resemblances, accord-
ing as they may indicate a probable relationship with plants or
animals, serve as a guide in determining the position of an
organism.
While itthus impossible to distinguish sharply the two great
is

groups of living organisms from one another, a distinction between

them and lifeless bodies is readily recognised. Living organisms are
endowed with the quality of IRRITABILITY, in which all lifeless
bodies are deficient. External or internal stimuli influence living
organisms to an activity, which is manifested in accordance with
the requirements and conditions of their internal structure. Even
in the smallest known organisms the manifestations of life are
occasioned by a similar sensitiveness to external or internal stimuli.
It is, therefore, probable that the simplest living beings must have
possessed simpler properties than any organisms now
essentially
known, which would enable us to connect them with non-living
substances. The substance which serves as a basis for all develop-
ment must be supposed to have had an inorganic origin. So far as i-
actually known, however, all living organisms have arisen only from
similar organisms. So far as experience has shown, spontaneous
generation is unknown. In the olden times it was a common
supposition, which ARISTOTLE himself held, that even highly organised
animals and plants could originate from sand and mud. In the
same degree that knowledge of the actual development of living
organisms was extended, the previously accepted cases of spontaneous
generation became more and more restricted, and were finally
limited to intestinal worms which could not
otherwise, it was
thought, be accounted for, and to microscopic organisms, the origin
of which also was not understood. Now, for such organisms the
 INTRODUCTION

possibility of aspontaneous generation has been disproved by more

modern investigations the history of the development of intestinal
;

worms is known, and the germs of minute organisms have been

found to exist everywhere. SCHWANN and PASTEUR have been
pioneers in this work, and have shown that it is possible to hinder
the development of the lower organisms, in places where it is
customary to find them, by destroying all existing germs and at
the same time preventing the entrance of new ones. It is due to
the results obtained by these men in their investigations on
spontaneous generation that we are now able to preserve food in
a scientific manner. The germs previously existing in the substance
to be conserved are destroyed by heat, while, by a proper mode of
sealing, the entrance of new germs is rendered impossible, and the
decomposition, which their presence would occasion, is accordingly
prevented.
All known living organisms have been derived from other
living organisms. But the idea of the origin of living from dead
substances has on the other hand derived important support from
the progress of chemical research. In the early decades of the
last century it was customary to draw a distinct line of separation
between organic and inorganic chemistry, and to assume that the
substances dealt with by organic chemistry could only be produced
by the vital action of organisms. The laws governing inorganic
chemistry appeared to have no reference to organic chemistry, the
formation of organic substance being due to a special force, the
li
vital force." In 1828 WOHLER obtained urea from ammonium
cyanate, and thus for the first time produced an organic compound
from an inorganic substance. In 1845 KOLBE completely synthesised
trichloracetic and in 1850 BERTHKLOT synthesised alcohol
acid,
and formic The former substance had been synthetically
acid.

prepared by HENNEL in 1828, but BERTHELOT was the first to

recognise its identity with the substance formed in alcoholic
fermentation. By these results the former distinction between
organic and inorganic chemistry was destroyed. Organic chemistry
has become the chemistry of carbon compounds.
In some such way it is possible that living matter originated from
non-living at some period in the evolution of the earth when the
conditions for its formation existed. In order that the organic
world should have developed from the first living matter, one of
the original properties of the latter must have been a capability of
continued existence among its surroundings. It must have been

capable of variation and of retaining the new characters appearing

in this way, of growth, i.e. the increase of itself at the cost of
foreign substances, and of reproduction, multiplication by
i.e.

separation into a number of parts. Some observers have recently

described the origin of microscopic structures which behave similarly
6 BOTA X Y

to living beings in bouillon anil other organic culture^ media when

7
exposed to the action of radium ( ).
Botany, or the science of plants, may be divided into a general
and a special part. In the general part, the structure and functions
of plants as such will be considered ; in the special part, the
particular structure and functions of the separate orders of plants
will l)e discussed.
The study of the structure of plants is called MORPHOLOGY ;

that of their functions PHYSIOLOGY. In the general part, morphology

and physiology will be treated separately; in the special part,
conjointly.
 PART I

GENERAL BOTANY

SECTION I

MOEPHOLQGY
 GENERAL BOTANY

SECTION 1

MORPHOLOGY
Tin; object of vegetable morphology is the scientific study of the
forms of plants. It does not attempt to discover the causes of the
variation in the forms, but rather has accomplished its purpose when
it succeeds in
showing how one form may be derived from another.
The only real basis of morphological study is, accordingly, the genea-
logicaldevelopment or phylogeny (p. 2). As phylogenetic develop-
ment can only be inferred, and cannot be directly followed, the
methods of morphology must also be indirect. They are dependent
on the one hand upon ontogeny, i.e. on the study of the development
passed through by an organism in attaining its mature condition,
and on the other hand upon the comparison of existing organisms
with one another and with those that have become extinct. To a
certain extent the ontogenetic development of a plant repeats its
phylogeny and helps to elucidate the latter, while, by means of com-
parative investigation, extreme forms may be connected by inter-
mediate links. As, however, the ontogeny of a plant is neither a
complete nor invariable repetition of its phylogeny, and as connecting
links between extreme forms are often wanting, the results of
morphological study are frequently incomplete. Such parts or members
of plants which it reasonable to presume have had a common
is

origin are distinguished as HOMOLOGOUS ; those which, while probably

having different origins, yet exercise the same functions, are termed
ANALOGOUS. Through the adaptation of different parts to the same
function, a similarity in both external form and internal structure
often results ;
and in this way the correct determination of morpho-
logical relationships is rendered difficult. Only homologous parts
have the same " morphological value." This homology is determined
10 BOTANY PARTI

by the facts of phylogeny and origin, and not by any correspondence

in function. Though the function of any structure does not influence
its morphological value, the need of making clear the intimate con-
nection between form and function often introduces physiological
considerations into morphological questions. When, for phylogenetic
reasons, it seems possible to attribute to a number of different
members a common origin, such a hypothetical original form is
termed the fundamental or primitive form. The various modi-
fications which the primitive form has passed through constitute its
METAMORPHOSIS. In this way the theory of the metamorphosis of
plants,which was once but an ideal conception, acquires an actual
significance.
Slightly differentiated structures, which are found at the beginning
of a series of progressively differentiating forms, are termed i;n>i
MENTARY ; imperfect structures, which have arisen as the result of the
deterioration of more perfect forms, are termed REDUCED.
Vegetable morphology includes the study of the external form and
the internal structure of plants. The descriptive study of the external
form of plants has been termed organography ( 8 ). This term will
not be used, since by the use of the word " organ," it would seem to
have a physiological signification. Morphology takes no recognition
of the parts of a plant as organs, but treats of them merely as
members of the plant body. On the other hand, one of the most
important aims of physiology is to place the external form and the
internal structure of the living body in relation to the functions
performed by the latter ; physiology also investigates the causes of
the organisation. The study of the internal structure of plants is
often designated Anatomy or Phytotomy ; but as it usually includes
also the study of the more minute internal structure, it resembles
rather histology, in the sense in which that term is used by zoologists,
and concerns itself to a much less degree with the coarse anatomy of
the plant body. In any case, it is the simplest plan to designate the
study of the outer form EXTERNAL MORPHOLOGY, and that of the
inner structure INTERNAL MORPHOLOGY.

I. EXTERNAL MORPHOLOGY (
9
)

Plants show a great diversity in the form and arrangement of

their members it is the task of morphology to determine the points
;

of agreement existing between them. It seeks to do this by deter-

mining the common origin of the homologous parts or members.
SECT. I MORPHOLOGY 11

The Development of Form in the Plant Kingdom

The Thallus ( ). When the body of a plant is not differentiated
10

into separate members, or is composed of members which (though they

may be similar) are not
homologous with those of
the most highly organised
plants, it is termed a
2 ---s
THALLUS. When the
. ,.. , .
Fl -

txrevisiae.
'""
*?
Cells
,, ,, 1,
thallus is differentiated wlthout bu ds ;
-i and
into members analogous 3, budding ceils, (x
540-)
to those of
higher the
plants some confusion may arise from the
Fm.i.-Gtottifa At
same names being used for parts which,
Commencement of division ; B, since their origin has been distinct, are not
(to the left) shortly aftendi vision ;
homoloo-QUSS-
C', a resting stage, (x MO.) .

simplest form that we can imagine

The
for an organism is that of a sphere, and this is actually the form of
some of the lower plants. A green growth often seen on damp

Km. 4. Bacteria from deposits on teeth, c,

Leptothi-ii' hum i//'x; "', the same after treat-
ment with iodine; b, Micrococcus c, Spin- ;

chaete dentium after treatment with iodine ;

il. *pii-!Uit,ii .tntitiiji-rtiim. (X 800.)

FIG. 3. Pinnului-iu viridi*. A,

view ; B, lateral view. ( x 540.)

walls consists of an aggregation of the microscopically small spherical

bodies of Gloeocapsa poli/dermatica (Fig. 1), an Alga belonging to one of
12 IJOTAXY

the lowest divisions of the vegetable kingdom. The single plants of

the Beer-yeast (Saccharomyces cererisiae) are ellipsoidal but, from their ;

peculiar manner of growth, by budding, they form lateral

outgrowths, and thus often appear constricted (Fig. 2).

Cylindrical and also disc -shaped forms are shown by
various Algae. The Diatomeae (Fig. 3), in particular,
exhibit a great variety of spindle, canoe, helmet, and
fan-like shapes ; but they may all be derived from the
more simple spherical, discoidal, or cylindrical forms.

1' ".. Plea Lactuca.

young stage, show-
ing apex anil base.
x 220.)

Fio. 0. Portion of Cluiln/,: Flu. ~. <'lail"*tiiJins ivrtii-illatn.". (After

(X 48.) TRINOSHETM, x 30.)

Among the Bacteria, which, as the cause of infectious diseases and

of decomposition, have been the object of so much recent investiga-
tion, we also meet with spherical, rod-shaped, filamentous, and
spirally wound forms (Fig. 4). The next stage in the progressive
SECT. I MORPHOLOGY

development of external form in the vegetable kingdom is exhibited

by such plants as show n DIFFERENTIATION INTO APEX AND P.ASE.
The base serves as a point
of attachment, while growth
is localised at the apex. In
this a growing point is
way
developed at the apex. As
an example of such a form,
a young plant of the green
Alga, Ufoa Laduca (Fig. 5),
may be taken. The de-
velopment of a more com-
plicated external form is
represented by the branched
filamentous, or ribbon-
Fio. 8. Dictyota dichotnmn. (jj nut. size.)
shaped Algae, in which the
origin of new formations is more and more restricted to the
apex.
An ACROPETAL order of development, in which the youngest lateral
members are always nearest the
growing apex, is clearly demon-
strated by the branched fila-
ments of the common green
Alga, Cladophora glomerata (Fig.
6). Still more pronounced is

the apical growth in the brown

seaweed Cladostephus rerticillatus
(Fig. 7). The great variety
in the form of the larger Fungi
and Lichens, by which they
are distinguished as club-, um-
brella-, salver-, or bowl-shaped,
or as bearded or shrub -like,
comes about by the union or
intertwining of apically grow-
ing filaments. This type of
construction is limited to Fungi
and Lichens. As the apex itself
may undergo successive bifurca-
tion, as in the case of Dictyota
dichotoma (Fig. 8), it does not
always necessarily follow that
FlO. 9. Hydi-<ili'ji"tli'nii (o nut. sixr.) new members must be formed
beneath the original apex.
The highest degree of external among the lower
differentiation

plants is met with in certain groups of red and brown sea-weeds

(Rhodophyceae and Phaeophyceae). Many representatives of these
14 BOTANY

classes resemble the higher plants in the formation and arrangement

of their members; Hydrolapathum sant/uim-iim (Fig. 9), for example,
as is indicated by its name, resembles a species of fittmex, and
affords an instructive illustration of the analogy of form existing

Fia. 10. Kiccin ii<iitmi.<

(Nat. slice.')

Fio. 11. Blasia piisillu. s, Sporogoiiiuin ;

r, rhizoids. (x 2.)

between plants phylogenetically widely distinct from one another.

The progressive differentiation of the thallus of the Bryophyta
(Mosses and Liverworts)" presents an even closer parallel to the
organisation of the higher plants. Within
the group of the Liverworts (Hepaticae) are
found ribbon-shaped forms and others that
show a division into distinct members.
Thus the thallus of fiiccia jlnitans (Fig.
10) is ribbon -shaped and dichotomously
branched, and its habit or general ap-
pearance recalls the Brown Alga Didyota
ilichotoma mentioned above (Fig. 8). Blasia

pusilla(Fig. 11) has marginal indentations

in its ribbon-shaped body. Lastly, Plagio-
chila asplenioides (Fig. 12), another Liver-

wort, has a distinction of stem-like and

leaf-like members which is completely
analogous to that exhibited by the most
highly organised plants. In spite of this
variety in external form, and the high
organisation that may be reached, the
Ki<:. 1-.- -I'luiiiiK-Iiilit ax/ill

.--.
S[>orogoiiiuin. (Xat. size.) body of these lower plants is termed a
thallus, and they are grouped together as
Thallophytes in contrast to the higher plants or Cormophytes.
The Cormus. All plants, from the Pteridophtya or fern-like
plants onwards, may be grouped together as CORMOPHYTES.
It

may be assumed that they have had a common origin, and that
 MORPHOLOGY 15

the similarity of their organisation is due to true homology. In

addition to the distinction of stem and leaf in their shoots, they possess
true roots, while even the most highly organised Bryophyta have
only filamentous structures (rhizoids) (Fig. 11 r) in place of roots
to attach them to the substratum. True roots, on the other hand,
which appear for the first time in the Pteridophyta, are, for the most
part, cylindrical structures with apical growth. Besides possessing
a distinctive internal construction they are distinguished in their
external form from the shoot by having a special sheath, the ROOT-
CAP or CALYPTRA covering the growing point, and by the absence
of leaves. The body of the higher plants provided with stem,
root, and leaf is termed a cormus, and the plants are spoken of as
cormophytes.
The Metamorphosis of the Primary Members of Cormophytes.
After the differentiation into stem and leaf and the appearance of
the roots had taken place, further changes have consisted essentially
in a more or less profound modification of these primary members
of the cormophytic plant-body. Such changes are spoken of as a
metamorphosis (p. 10), and in some cases may be so extensive as to
lead to one primary member assuming the characters of another.

The relationships between homologous members, which are often vei'y striking,
did not escape the notice of earlier observers. They suggested comparisons,
although no real phylogenetie basis for such comparisons existed. Thus, an ideal-
istic conception of the form of external members was developed, and finally reached
its highest artificial development in GOETHE'S Theory of Metamorphosis ;
and its
scientific conclusion in the writings of ALEXANDER BIIAUN. As the great variety
exhibited in the external appearance of the lower plants precluded any possibility
of assigning to them hypothetical primitive forms, the whole terminology of the
external morphology of plants has been derived from conceptions applicable only
to the Cormophytes. Even to-day, the same terms used in reference to the
Cormophytes are applied to parts of the Thallophytes, which are evidently only
analogous.

Members of Independent Origin. Parts which cannot be derived

by metamorphosis of the
primary members of the cormophytic plant
are sometimes met with. Though they are of infrequent occurrence
they are of importance as showing that the natural evolutionary
process is not to be limited by any formal scheme.. Such structures
will be discussed farther on.

Relations of Symmetry
Every section through a part of a plant, made in the direction of
its longitudinal axis, is distinguished as a longitudinal section those
;

at right angles to it being termed cross or transverse sections.

Parts of plants which may be divided by a number of longitudinal
planes into like halves are termed either MULTILATERAL, RADIAL, or
ACTINOMORPHIC. Such parts are symmetrically constructed around
16 BOTANY

their longitudinal axis. The degree of symmetry peculiar to any

leafy shoot will be more apparent from a diagram, that is if the

KIC;. lli. -Diagram showing tin- su-callfl cl> . 14. Diagram showing two-tmoked
rnssiiti- arrangement <>t' leaves. alternate arrangement of lea\

leaves which it bears be projected on a plane at right angles to its

axis. The radial symmetry of a shoot with opposite leaves is clearly
shown in the adjoining diagram (Fig. 13).
I
lV A shoot with its leaves arranged alternately
in two rows shows somewhat different rela-
tions of symmetry. The diagram of such
a shoot (Fig. 14) can only be divided into
similar halves by two planes. When such
a condition exists, a member or plant is
said to be BILATERAL. When, however, a
division into two similar halves is only
possible in one plane, the degree of sym-
metry is indicated by the terms DORSI-
VENTRAL or since, while the
ZYGOMORPHIC;
right and halves correspond to one
left

another, differences exist between the dorsal

and ventral surfaces. Ordinary foliage-
leaves exhibit this dorsiventral structure.
In the accompanying figure (Fig. 15) such
a
monosymmetrical, dorsiventral foliage-
diagrammatical ly represented. From
leaf is
the surface view (A) and from the cross-
.v
section (/?), in which the distinction between
15.
<i. Diagram of a foliage-leal. the dorsal and ventral sides is indicated
.1, Surface view trausverse ; li,
by
section plane of symmetry.
: -. shading, it is obvious that but one plane of
symmetry (s) can be drawn. Dorsiventral
members are often ASYMMETRICAL, not being divided by any plane
into corresponding halves : the leaves of many kinds of Begonia will
SKUT. I MORPHOLOGY 17

serve as examples of this. In such cases and the leaf of the Elm
may be mentioned as another striking example the symmetry of
the individual leaf is subordinated to that of the entire plant.

Branch Systems

Thallophytes as well as Cormophytes exhibit systems of branching,

resulting either from the formation of new growing points by the
bifurcation of a previously existing growing point, or from the develop-

Ma
Fie. 10. Diagrams of branch systems. A, Dichotoinous branching Aa, ci|iial dichotomy Ab,
; ;

scorpioid dichotomy Ac, helicoid dichotomy, li, monopodial branching lla false dichotomy ;
; ;

Kb, scorpioid cyme Be, helicoid cyme ., s, sympodia.

; ;

ment new growing points in addition to those already present.

of
In way there arise two systems of branching, the DiCHOTO-
this
MOUS and the MONOPODIAL. By the uniform development of a
continuously bifurcating stem, a typical dichotomous system of
branching is produced, such as is shown in Dictyota dichotoma (Fig. 8),
and is represented diagrammatically in Fig. 16 Ad. In a typically
developed example of the monopodial system there may always be
distinguished a persisting main axis, the MONOPODIUM, giving rise
to lateral branches from which, in turn, other lateral branches are
developed. A
good example of this form of branching is afforded
by a Fir-tree. Where one of the two branches is regularly developed
at the expense of the other, the dichotomous system assumes an
c
18 BOTANY PART I

appearance quite different from its typical form. The more vigorous
branches may then, apparently, form a main axis, from which the
weaker branches seem to spring, just as if they were lateral branches.
This mode of branching (Fig. 16 Ab} is illustrated by the Selaginellas.
Such an apparent main axis (s, s) is termed, in accordance with its
origin, a SYMPODIUM. On
the other hand, in the monopodial system
two or even several lateral branches may develop more strongly than
the main axis, and so simulate true DICHOTOMY or POLYTOMY. Such
monopodial forms of branching are referred to as FALSE DICHOTOMY
(Fig. 16 Ea) or FALSE POLYTOMY, as the case may be. A
good
example of false dichotomy may be seen in the Mistletoe (Viscum
itHniin). If, however, a lateral branch so exceeds the main axis in

development pushing the apex of the latter to one side, that it seems
ultimately to become a prolongation of the axis itself, a sympodium is
again formed (Fig. 16 Eh). This is what occurs in many of our forest
trees, e.g. the Lime and Beech ; in both of these trees the terminal buds
of each year's growth die, and the prolongation of the stem, in the
following spring, is continued by a strong lateral bud, so that in a
short time its sympodial origin is no longer recognisable. In many
rhizomes, on the other hand, the sympodial nature of the axis can be
.easily distinguished ; as, for example, in the rhizome of Polygonatum
multiftorum (Fig. 23), in which, every year, the terminal bud gives
rise to an aerial shoot, while an axillary bud provides for the
continuance of the axis of the rhizome. In the flower-producing
shoots or inflorescences of Phanerogams the different systems of
branching assume very numerous forms. These will be more fully
described in their proper place. To such inflorescences belong the
ventrally coiled dorsiventral shoots which produce new shoots from
their convex dorsal surfaces, instead of in their leaf -axils.

The Shoot

The Development of the Shoot. Under the term shoot a stem

and its leaves are collectively included. A stem possesses an apical
mode of growth (Fig. 17), and its unprotected growing point is
described as naked, in contrast to that of the root with its sheathing
root-cap. The apex of the shoot generally terminates in a conical
protuberance, called the VEGETATIVE CONE. As it is usually too
small to be clearly visible to the unaided eye, it is best seen in magnified
median longitudinal sections. So long as the apex of the shoot is
still internally undifferentiated, it continues in the embryonic con-

dition, and it is from the still embryonal vegetative cone that the
leaves take their origin. They first appear in acropetal succession
as small, conical protuberances, and attain a larger size the farther
removed they are from the apex of the stem. As the leaves usually
grow more rapidly than the stem which produces them, they envelop
 MORPHOLOGY 19

the move rudimentary leaves, and, overarching the vegetative cone,

form a BUD. Buds are therefore merely undeveloped shoots. If they
are to remain for a long time undeveloped, as for example is the

FH:. 18. Longitudinal section f :i

bifurcating shoot (;>) of I./ico-
podlum alpinimi, showing equal
development of the rudimentary
PIG. 17. Apex of u. shoot of a phanerogamic shoots, p', p" ; b, leaf rudiments ;

plant, Vegetative cone /, leaf rudiment

v, ; ; c, cortex; /, vascular stninds.
g, rudiment of an axillary bud. (x 10.) (After HF.UKI.MAIKK, x i>0.)

case with winter buds, they are protected in a special manner during
their period of rest.
The Origin of New Shoots. The formation of new growing
points by the bifurcation of an older growing point, in a manner
similar to that already described for Dictyota dichotoma (Fig. 8), occurs
also in the lower thalloid Hepaticae (Riccia
fluitans, Fig. 10). Among the Cormophytes
this method of producing new shoots is of
less frequent occurrence, and is then mainly
limited to the Pteridophytes, and is typically
shown only in some Lycopodiaceae. In this case,
whenever a shoot is in process of bifurcation, two
new' vegetative cones are formed by the division FIG. 19. Hifurcating shoot
(p) of Lycofniilhitii hin i"
of the growing point (Fig. 18). In most of the datum, showing unequal
Lycopodiaceae the new shoots thus formed de- development of the rudi-
the weaker becomes pushed mentary shoots, ]>', p" ;

velop unequally ;
b, leaf rudiments. (After
to one side and ultimately appears as a lateral HEGKLMAIF.R, x 40.)
branch (Fig. 19). Although a relationship as re-
gards position is generally apparent between the origin of leaves and
the lateral shoots, in the system of branching resulting from such a
bifurcation of the vegetative cone this connection does not exist.
In the more highly developed Bryophytes, particularly in the true
Mosses, new shoots arise obliquely below the still rudimentary leaves
20 BOTANY PART i

at some distance from the growing point. In the Phanerogams new

shoots generally arise in the axils of the leaves. In the accom-
panying illustration of a longitudinal section of a phanerogamic
shoot (Fig. 17) the rudiment of a shoot (g) is just appearing in
the axil of the third uppermost leaf in the axils of the next older
;

leaves the conical protuberances of the embryonic leaves are

already beginning to appear on the still rudimentary shoot. Shoots
thus produced in the axils of leaves are termed AXILLARY SHOOTS.
The leaf, in the axil of which a shoot develops, is called its SUH-
TENDING LEAF. An axillary shoot is usually situated in a line
with the middle of its subtending leaf, although it sometimes
becomes pushed to one side. As a rule, only one shoot develops
in the axil of a leaf, yet there are instances where it is followed

by additional or ACCESSORY SHOOTS, which either stand over one

another (serial buds), as in Lonicera, Gleditschia, Gymnoclcuhis, or side
by side (collateral buds), as in many Liliaceae, e.g. species of AUnnn
and Muscari.
Although in the vegetative regions, i.e. the regions in which
merely vegetative organs are produced, the rudiments of the new
shoots of phanerogamic plants make their appearance much later
than those of the leaves, in the generative or flower-producing regions
the formation of the shoots follows directly upon that of their sub-
tending leaves, or the shoots may even precede the leaves. In this
last case the subtending leaves are usually either poorly developed
or completely suppressed, as in the inflorescence of the Cruciferae.
The bud forming the end of a shoot is called the terminal bud,
while those borne on the sides of the shoot are the lateral buds.
Shoots developing in predetermined positions on young parts
of the plant are designated NORMAL, in contrast to ADVENTITIOUS
SHOOTS, which are produced irregularly from the old or young portions
of a plant. Such adventitious shoots frequently spring from old
stems, also from the roots of herbaceous plants (Brassica oleracea.
Anemone sylvestris, Convolvulus arrensis, Eumex Acetosella), or of bushes
(J,'n!iiis, Rosa, Corylus), or
of trees (Populus, Ulmus, Rolinia). The)'
may even develop from leaves, as in Cardamim pratensis, Nasturtium
and a number of Ferns. An injury to a plant will frequently
officinale,
induce the formation of adventitious shoots, and they frequently arise
from the cut surface of stumps of trees. Gardeners often make use
of pieces of stems, rhizomes, or even leaves as cuttings from which
to produce new plants.
Leaves and also normal shoots, which make their appearance as out-
growths from the portions of the parent shoot, still in an embryonic
condition, have an external or EXOGENOUS origin. Adventitious
shoots, on the other hand, which arise from the older parts of stems
or roots, are almost always ENDOGENOUS. They must penetrate the
outer portions of their parent shoot before becoming visible. Adven-
SECT, i MORPHOLOGY 21

titious shoots formed on leaves, however, arise, like normal shoots,

exogenously.

Buds are formed in the marginal indentations of the fleshy leaves of species
of Bryophyllum (Crassulaceae). Although arising from the leaf these buds
must properly be regarded as "normal," and as forming part of the normal
ontogeny of the plant, since they arise in pre-determined positions from young
tissue. In the strict sense of the term only those buds can be called adventitious
which are produced in casual positions from tissues which in their production
enter into renewed activity, e.g. the buds which arise at the base of isolated leaves
of Beyonia when these are laid on damp soil. The concept of " normal b'uds" is
notwithstanding usually taken in a narrower sense and limited to buds which
arise on the axis of the shoot in the normal course of development. Examples,
however, occur which make it difficult to draw a sharp distinction in this latter
sense between normal and adventitious shoot-formation ( n ).
While, as a rule, new leaves arise beneath the apex of a stem, which
continues to grow, exceptions to this occur, especially in floral structures the ;

apical cone may be used up in the formation of a terminal leaf-rudiment. In this

way many of the stamens and carpels of Phanerogams, which are described as
terminal, arise.
In some shoots of limited growth such as the inflorescences of many
Phanerogams (Vallisneria) new members arise in an inverted order
lateral

proceeding from the apex towards the base. The introduction of new lateral
members between those already present is also met with in the floral region ( 12 ).
It was mentioned above that normal shoots arose from the embryonic tissue
of the growing point of the parent shoot. When they are apparent at a greater
distance from the apex (Fig. 17) it can usually be shown that embryonic substance
has been reserved at the proper points for their formation. The growing points of
adventitious shoots are for the most part derived from embryonic tissues which
have persisted iu the older portions of the plant and are capable of increase.
They can, however, also arise from older tissues, owing to the capability of the
latter to return to the embryonic condition and produce new growing points.

Further Development of the Shoot. The processes of develop-

ment, which result in the production of new members at the apex of
a shoot, are followed by their increase in size and further growth.
This growth is usually introduced by the vigorous elongation of
the segments, which is spoken of as growth in length, while at the
same time the buds expand. The zone of most rapid growth in
length is at some distance from the growing point. The growth
in length and consequent elongation of the shoot is in some cases
so slight that the leaves remain close together, and leave no free
spaces on the stem, thus forming so-called SHORT SHOOTS. The
same plant may bear ordinary elongated shoots and short shoots.
The duration of life of the latter is usually shorter, and they fre-
quently do not branch, and take no part in the permanent branch
system of the tree. This is seen in the Larch, in which the
short shoots with crowded needle-shaped leaves are borne on
elongated shoots. The stem of a shoot, as contrasted with the leaves,
22 BOTANY PART I

isoften spoken of as the AXIS the portions of the stem axis between
;

the insertions of the leaves are termed the INTERNODES, and the parts
of the axis, from which the leaves arise, the
NODES. When the base of the leaf encircles
the stem, or when several leaves take their
origin at the same node, the nodes often be-
come strongly marked (Labiatae).
In some cases the growth in length of a
shoot continues for a longer time at certain
intermediate points by means of INTERCALARY
<;ROYVTH. Such regions of intercalary growth
are generally situated at the base of the inter-
nodes, as in the case of the Grasses. dis- A
placement from the position originally occupied
by the members of a shoot frequently results
from intercalary growth. A bud may thus,
for example, become pushed out of the axil
Kic;. -JO. Xaiiiolus I'/'Icraiidi, . ., Ti , e j
i A i i_

each shoot of lts subtending leaf, and so apparently have

axillary (a) .

bearing its subtending leaf its

origin higher on the stem; or a subtending
1 8 '" *
1 '
leaf in the COU1>Se * itS ma J Can7 its
fruit*"' iSt'si""" '
growth*
axillary bud along with it, so that the shoot
which afterwards develops seems to spring directly from its subtend-
ing leaf or, finally, the subtending leaf may
;

become attached to its axillary shoot, and

growing out with it, may thus appear to spring
I.- tin
from it (Fig. 20).
Iu the annual growth in length of a shoot of a tree,
i.e. the increment formed during one vegetative period,
tlio loAvest internodes which intervene between the bud-

scales are very short. By means of the closely crowded

scars of the bud-scales the limits of the shoots formed in
successive years can be determined.

Resting Buds. As a means of protection

against loss of water in our latitudes, the buds
of shrubs and trees are usually invested, in
winter, with scale-like leaves or BUD-SCAL1>
(Fig. 21). These are rendered still more effec-
tive as protective structures by hairy outgrowths
and excretions of resin and gum, and also by
the occurrence of spaces filled with air between
the scales. Not infrequently
^ J the subtending & ,
Fin. 21. Winter bu<is of the
leaf takes part in the protection of its axillary Beech (*,.,"'>

bud, the base of the leaf-stalk, after the leaf *. Bud--to* (Nat.

itself has fallen, remaining on the shoot as a

cap-like covering for the winter bud. The buds of tropical plants,
which have to withstand a dry period, are similarly protected ; but
 MORPHOLOGY 23

where the rainfall is evenly distributed throughout the year buds

develop no such means of protection.
In many deciduous trees, such as the Willow, the terminal buds of the year's
growth regularly die. In nearly all trees many buds, iisually the first-formed
buds of each year's shoot, seem able to remain dormant during many years without
losing their vitality these are termed DORMANT BUDS.
: In the case of the Oak or
Beech such latent buds can endure for hundreds of years in the meantime, by ;

the elongation of their connection with the stem, they continue on its surface.
Often it is these, rather than adventitious buds, which give rise to the new growths
formed on older parts of stems. It may sometimes happen that the latent buds
lose their connection with the woody parts of their parent stem, but nevertheless

grow in thickness, and develop their own wood they then form remarkable
;

spherical growths within the bark, which may attain the size of a hen's egg and
can be easily separated from the surrounding bark. Such globular shoots are
frequently found in Beech and Olive trees.

i,
FIG. 23. Rhizome of Polygonatum tnultiflorum. a, Bud of
next year's aerial shoot ; b, scar of this year's, and
c, d, e, scars of three preceding years' aerial shoots
; ,

roots. (^ nat. size.)

FHJ. 22. Shoot of Dentaria bulbi

fera, bearing bulbils, br. ,
(Nat.
size.)

The Metamorphosis of the Bud. The BULBILS and GEMLE>

which become separated from the parent plant and serve as a means
of reproduction, are special forms of modified buds. To facilitate
the storage of reserve nutritive substances they have usually the form
of small tuberous bodies. Many plants owe their specific name to
the that they produce such bulbils, as, for example, Lilium
fact

bulbiferum and Dentaria bulbifera (Fig. 22).

The Metamorphosis of Subterranean Shoots. Shoots that live
underground undergo characteristic modifications, and are then termed
24 BOTANY PART I

ROOT-STOCKS or RHIZOMES. By means of such subterranean shoots

many herbaceous perennial plants are enabled to persist through the
winter. A rhizome develops only reduced leaves in the form of
larger or smaller, sometimes scarcely visible, scales. By the presence
of such scale leaves, with their axillary buds, and by the absence of
a root-cap, as well as by its internal structure, a rhizome may be dis-
tinguished from a root. Rhizomes usually produce numerous roots ;
but in a few cases these are wanting and the rhizome itself functions
as a root. Rhizomes often attain a considerable thickness and store

zk

Fio. 24. Rhizome of Coralllorrhiza in- FIG. 25. Longitudinal section of Tulip
nata. a, Floral shoot ; 6, rudiments bulb, Ttdipa Gesnerwna. zk, Muililioi
of new rhizome branches. (After stem ;i zs, scale leaves ; v, terminal
SHACHT, nat. size.) bud ; k, rudimentof a younj; bulb ; ,

roots. (Nat. si/c.)

up material for the formation of aerial shoots.

nutritive In the
on the preceding page (Fig. 23) is shown the root-stock of
illustration
Solomon's Seal (Polygonatum multiflorum). At d and c are seen the
scars of the aerial shoots of the two preceding years ; and at b may
be seen the base of the stem growing at the time the rhizome was
taken from the ground, while at a is shown the bud of the next
year's aerial growth. The rhizome of Coralliorrhiza innata, a sapro-
phytic Orchid, which grows in soil rich in humus, affords a good
example of a root-stock functioning as a root (Fig. 24). BULBS, also,
belong to the class of metamorphosed shoots. They represent a
shortened shoot with a flattened, discoid stem (Fig. 25 zk), the fleshy
 MORPHOLOGY 25

thickened scale (zs) of which are filled with reserve food

leaves
material. The growth of a bulb develops from its axis, while
aerial
new bulbs are formed from buds (k) in the axils of the scale leaves.
Another form of underground shoot, allied to bulbs and connected
with them by transitional forms, is distinguished as a TUBER. The
axis of a typical tuber, in contrast to that of a bulb, is fleshy and
swollen, serving as a reservoir of reserve material, while the leaves
are thin and scaly. Of such tubers those of the Meadow Saffron
(Colchicum autumnale) or of Crocus sativus are good examples. In the
Meadow Saffron new tubers arise from axillary buds near the base of
the modified shoot, but in the Crocus from buds near the apex. In
consequence of this, in the one case the new tubers appear to grow
out of the side, and in the other to spring from the top of the old
tubers. The tubers of the Potato (Fig. 26) or of the Jerusalem
Artichoke (Helianthus tuberosus) are also subterranean shoots with
swollen axis and reduced leaves. They are formed from the ends of
branched, underground shoots or runners (STOLONS) and thus develop
at a little distance from the parent plant. The so-called eyes on the
outside of a potato, from which the next year's growth arises, are in
reality axillary buds, but the scales which represent their subtending
leaves can only be distinguished on very young tubers. The parent
plant dies after the formation of the tubers, and the reserve food
stored in the tubers nourishes the shoots which afterwards develop
from the eyes. Since, in their uncultivated state, the tubers of the
Potato plant remain in the ground and give rise to a large number
of new plants, it is of great advantage to the new generation that the
tubers are produced at the ends of runners, and are thus separated
from one another.
The Metamorphosis of Aerial Shoots. Similar advantages to
those obtained by the elongation of the underground shoots in the
Potato accrue from surface runners, such as are produced on Straw-
berry plants. Surface runners also bear scale-like leaves with axillary
buds, while roots are developed from the nodes. The new plantlets,
which arise from the axillary buds, ultimately form independent
plants by the death of the intervening portions of the runners.
Still more marked is the modification exhibited by shoots
which only develop reduced leaves, while the stems become flat and
leaf-like and assume the functions of leaves. Such leaf-like shoots
are called CLADODES or PHYLLOC LADES, and Goebel proposes to dis-
tinguish those flattened shoots which have limited growth and specially
leaf-like appearance as phylloclades, and to term other flattened axes
cladodes. Instructive examples of such formations are furnished by
Ruscus aculeatus (Fig. 27), a small shrub whose stems bear in the
axils of their scale-like leaves (/) broad, sharp-pointed cladodes (cl),
which have altogether the appearance of leaves. The flowers arise
from the upper surface of these cladodes, in the axils of scale leaves.
26 BOTA N Y

In like manner the stems of the Opuntias (Fig. 28) are considerably

Fio. 20. Part of a growing Potato plant, Solatium tubero.tiiin. The whole plant lias
been developed from the dark-coloured tuber in the centre. (From nature,
copied from one of UAILLON'S illustrations, J nat. size.)

flattened, while the leaves are reduced to small thorny protuberances.

Fio. 27. Twig of Jtusciisaculeat'us. f, Leaf; show-in ,' llower

1

cladode flower.
I i' .
28. Opimtta MMOetMUta, Il,-iw.,
cl, ; bl, (Nat. size.) SCHUMANN.
and fruit. (After |.
nat. size.)

In this case the juicy flat shoots perform not only the functions of
SECT. I MORPHOLOGY 27

assimilatory organs, but also serve as water-reservoirs in time of

drought.
On the other hand a plant may lose its leaves more or less com-
pletely without any marked flattening or
thickening occurring in the stems, which
then take on a green colour this, for ex- ;

ample, is the case in the Broom (Spartium

scoparium), which-develops only a few quickly
falling leaves on its long, naked twigs. As
a rule, however, leafless green Phanerogams
will be found to have swollen stems, as in
the variously shaped species of Euphorbia
and Cactus.
Reduction of the Shoot in Parasites.
A great reduction in the leaves, and also in
the stems, often occurs in phanerogamic
parasites, in consequence of their parasitic
mode of life. The leaves of the Dodder
(Cusciita, Fig. 202
are only represented
b)
by very small yellowish scales, and the stem
is
similarly yellow instead of green. The
green colour would, in fact, be superfluous,
as the Dodder does not produce its own
nourishment, but derives it from its host
plant. Cuscuta Trifolii, one of the most fre-
quent of these parasites, is often the cause FIG. 2t>. Ampeiopsis veitckn.

Of the large yellow R R > < Stem-tendrils. (J nat.

f /I 1
sizp -)
areas frequently ob-
servable in the midst of clover fields. In
certain parasites belonging to the
tropical
family Rafflesiaceae, the process of reduction
has advanced so far that the flowers alone
are left to represent the whole plant. Eafflesia
is a re-
.l/'uoldi, a plant growing in Sumatra,
markable example of this its flowers, al-
;

though they are a metre wide, the largest

flowers in existence, spring directly from the
roots of another plant (species of Cisstis).
Tendrillar Shoots. A
peculiar form of
metamorphosis is exhibited by some climbing
FIG. 30. Stem-tin. in of GietHt- plants through the transformation of certain
schia triawnthos. (4 nat. of their shoots into TENDRILS. Such tendrils
size.)
assist the parent plant in climbing, either by

twining about a support or otherwise holding fast to it. The twining

bifurcated tendrils of the Grape-vine, for example, are modified shoots.
In some sub-species of the wild vine (Ampelopsis Jtederacea) and in
28 BOTANY PART i

other species of the genus, e.g. Am^dopsis Veitddi (Fig. 29), the
tendrils are able to form adhesive discs at the tips of the branches,
and thus to cling to flat supports.
Stem-thorns. Shoots may undergo a still greater reduction by
their modification into THORNS, as a defence against the depredations
of animals. Of shoots modified in this manner, the Black Thorn
(I'm a us spinosa), the White Thorn (Crataegus), and the Honey Locust
(Gleditschia) afford instructive examples. The thorns are simple or
branched, hard, pointed bodies. In Gleditschia (Fig. 30) the thorns
are developed primarily from the uppermost of several serial buds ;

while secondary thorns may develop on older portions of the stem

from the lower buds of the series, and thus give rise to clusters of
thorns. In Colletia cruciata all the shoots are flattened and spiny. ><>
that they perform the duties of the leaves, which are quickly lost, in
addition to serving as protective structures. This plant is an American
shrub belonging to the Rhamnaceae, and grows in dry sunny situations.
Flowers. The most marked changes in the form of the shoot, due
to the displacement and union of its different members, take place in
phanerogamic flowers (Fig. 39). The shoots from which flowers are
developed are termed FLORAL SHOOTS, in contrast to the FOLI.MJK
SHOOTS, the functions of which are merely vegetative. The axis of
the floral shoot remains short, and becomes flattened or even depressed
at the tip. The leaves which spring from the floral axis often become
united with one another and with the axis itself. In such cases
thorough investigation of the development and the comparative
morphology of the flower is necessary to reveal the modifications
which have taken place during its evolution. In most instances the
rule seems to hold that axillary buds are not formed within a flower
except in cases of abnormal development.
The Order of Sequence of Shoots. If the vegetative cone of the
primary axis of a plant, after reaching maturity, is capable of repro-
duction, a plant with but one axis will result, and the plant is
designated UNI AXIAL or HAPLOCAULESCENT. Usually, however, it is
not until a plant has acquired axes of a second or third order, when it
issaid to be DIPLOCAULESCENT or TRIPLOCAULESCENT, or of the nth
order, that the capacity for reproduction is attained. A
good illus-
tration of a plant with a single axis is afforded by the Poppy, in
which the firstshoot produced from the embryo terminates in a flower,
that is, in that organ of Phanerogams which includes the sexual
organs. As an example of a triplocaulescent plant may be cited the
common Plantain (Plantago major), whose primary axis produces only
foliage and scale leaves ; while the secondary axes give rise solely to
bracteal leaves, from the axils of which finally spring the axes of the
third order, which terminate in the flowers. In the case of trees,
only shoots of the nth order can produce flowers. Thus a division of
labour commonly occurs in a branched plant, which find-; its expression
 MORPHOLOGY 29

in differences of form between the successive shoots. These differ

in appearance according to the special function performed by them,
whether nutrition, storage, or reproduction. In addition to the
essential members in the succession of shoots developed in a deter-
mined order, there are non-essential members which repeat forms of
shoot already present. These may appear simultaneously with the
essential shoots, and serve to increase the size of the plant as in

many annuals ; in many perennial plants they arise as yearly innova-

tions on the stock. Adventitious shoots, as a rule, repeat members
which have already made their appearance. Exceptionally, they
form necessary links in the succession of shoots ; this is the case in
the Podostemaceae (an aquatic order of Dicotyledons found in the
tropics, the plants belonging to which resemble Lichens or Liverworts
in external form), where they bear the flowers.
The Habit or General Aspect of Plants is dependent upon the
origin, number, mode of growth, and duration of their branches, and
on the presence or absence of non-essential shoots. Cormophytes
which develop herbaceous aerial shoots, and persist only so long as
is requisite for the development and
ripening of their fruit, be it one
or several vegetative periods, are called HERBS. Herbaceous plants,
however, which, although annually dying down to the ground, renew
their existence each year by means of new shoots produced from

underground shoots, rhizomes, or roots, are further distinguished as

PERENNIALS or perennial herbs. SHRUBS or TREES, on the other
hand, have woody, persistent shoots, which bear fruit repeatedly.
In these the reproductive shoots are shed annually, and, in some,
vegetative branches are also cast off, the remaining ones persisting
and increasing in thickness. In the Lime the ends of the leafy twigs,
in the Scotch Fir the short shoots, and in the Oak, Elm, Willow, and

Poplar weak lateral branches are thus lost. The leaves of evergreen
trees remain alive for several years, while those of deciduous species

only persist for a single vegetative period.

Shrubs retain their lateral shoots, so that their branches are
formed near the ground trees, on the contrary, soon lose their lower
;

lateral branches, and have a main stem or trunk, which bears a crown
of branches and twigs. In many trees, shrubs, and herbs the main
shoot is vertical, while the lateral branches assume a horizontal
position, or are directed obliquely upwards or downwards. In other
cases the main axis is sympodial, a lateral branch continuing the
direction of growth of the primary shoot. Sometimes a main axis
is
indistinguishable among the group of similarly directed branches.
The general appearance of the plant is determined by the direction
and thickness of its branches and leafy twigs. If these are all
directed upwards the shape is pyramidal, while broadly pyramidal,
oval, and rounded forms arise when the branches diverge more
"
strongly. The " weeping varieties of several familiar trees are due
30 BOTANY PART i

to the branches becoming long and pendulous. Herbaceous plants

often have stems which creep on the surface of the ground. Other
plants, both herbaceous and woody, climb upon various supports by
means of prickles, and modified shoots, by means of
hooked hairs,
roots or tendrils, or by twining movements. Climbing plants are
termed LIANES, those which wind round a support being distinguished
as TWINING PLANTS. It is the presence of numerous rope-like stems
of lianes which renders the tropical forest so impenetrable. On both
creeping and climbing shoots the leaves tend to become displaced
towards the dorsal surface of the stem, while branches spring from
the sides and roots from the ventral surface.

In catalogues and descriptions of plants the duration of the period of growth

isusually expressed by special symbols : thus O
indicates an annual a biennial,
;

and H a perennial herb f? is employed to designate both trees and shrubs, and
;

for trees the sign fj is also in use.

Development of the Leaf. The first appearance of the leaf as a

lateral protuberance (Fig. 17 f) on the vegetative cone of the shoot
has already been referred to (p. 18). When the
apex of a shoot is removed by a transverse
section and viewed from above (Fig. 31), the
origin of leaves as lateral protuberances is more
evident than in a longitudinal section. The
embryonic leaf rudiment generally occupies
but a small portion of the periphery of the
f

vegetative cone ; it may, however, completely

surround it. In like manner, when the mature
leaves are arranged in whorls, the developing
protuberances of the rudimentary leaves may
form at first a continuous wall-like ring around

PIO. si. Apical view of thfi the growing point and only give rise later to
vegetative cone of a shoot the separate leaf rudiments. Leaves take their
of Eiwnvmus iavonicns. i e \ e i i

(x 12 )
origin only from such parts of a plant as have
remained in an embryonic condition. A leaf
never arises directly from the older parts of a plant. In cases where
it apparently does so its development has been preceded by the forma-

tion of a growing point of a new shoot. The growing point of a

shoot has usually 'an UNLIMITED GROWTH, while the growth of a leaf
isLIMITED. A leaf usually continues to grow at its apex for a short
time only, and then completes its segmentation and development by
intercalary growth, which is usually localised near the base. It is true
that some leaves, such as those of Ferns, have apical growth and
continue to grow for a long time (in some cases, e.g. Lygodium, the
growth is unlimited), and bear their pinnae in acropetal order. On
the other hand, the leaf-like cladodes, although they are in reality
SECT. I MORPHOLOGY 31

metamorphosed shoots, exhibit a limited apical growth like that of

ordinary leaves.
We have already seen that in certain cases the apical cone may give rise to a
terminal leaf. The apex of the leaf in many ferns (e.g. Adiantum Edgeworthii) may
on the other hand be transformed into a bud from which a shoot will develop ( 13 ).

The unsegmented protuberance of the still rudimentary leaf, termed

u
by EICHLER ( ) the primordial leaf (Fig. 32 A ft), first projects from
the vegetative cone of the
shoot (A v). This is usually
followed by a separation of
the primordial leaf into the
LEAF-BASE (g in A and B)
and the rudimentary lamina
or UPPER LEAF (o in A and
B). The leaf-base, or the
part of the rudimentary leaf
which immediately adjoins
Fl0 3 "- A i )ex of au Klm shoot Vl "<
'
j>erfri. A,
the vegetative cone, either -

Showing the vegetative cone r, with the rudiment!

'
. . , .
,
takes I1O further part in the of a young leaf, 6, still unsegmented, and of the next
older
leaf, exhibiting segmentation into the laminar
Succeeding differentiation of
thp Ipnf or ir dpvplnns into
intn rudiment, o, and leaf-base, g; showing the older
uie leai, o leveiops ]|laf viewed obliquely frolll behilld>
,

(x 58 }
a LEAF -SHEATH (vagina) or
into STIPULES. The upper leaf, on the other hand, gives rise to the
leaf-blade or LAMINA. If the fully developed leaf possesses a LEAF-
STALK (petiole), this is later interposed by intercalary growth
between the leaf-blade and the leaf-base.
The leaf-tip often develops more rapidly than the rest of the leaf, and GOEBEI.
regards this as an arrangement for the protection of the younger parts of the
bud ( ls ). This is seen most strikingly in some tropical plants, especially in climbers.
In this case, according to M. RACIBOKSKI ( I8 ), the "fore-runner tips" serve for
assimilation before the remainder of the leaf has attained the mature condition.
In leaves the laminae of which do not remain simple, but undergo segmenta-
tion as they develop, the lateral segments are as a rule formed in the basipetal
direction, i.e. from the tip towards the base (Fig. 33) ; the opposite direction of de-
velopment is, however, sometimes found. The segments of the palmate and pinnate
leaves of Palms originate by a process of splitting of the leaf blade which is, to begin
with, entire. The direction of the splits is determined by the folds of the lamina ( ).
17

Different forms of leaves are most clearly seen in the Phanerogams,

in which the various leaf structures found at different regions of the
shoot have been distinguished as SCALE LEAVES, FOLIAGE LEAVES,
BRACTEAL LEAVES, and FLORAL LEAVES (Fig. 34). These usually
succeed one another in definite order. However unlike the leaves
may become, their first origin -is similar. Since the scale leaves and
bracts can often be shown to be arrested forms of foliage leaves, the
latter may be first considered.
32 BOTANY

Foliage Leaves, generally referred to simply as leaves, are the

leaf structures on which devolves the task of providing nourish-
ment for the plant. Since the exercise of this function is dependent
upon the presence of the green pigment, foliage leaves have a
green colour. In certain cases, where their form is extremely

FIG. 33. Acer platanoides. A, external view of a bud, with two young leaves between
which the apical cone of the stem is visible sp, the leaf-blade, in which five
;

segments are indicated, the uppermost one being developed first st, the zone, by
;

the growth of which the leaf-stalk will arise later, li, an older leaf seen from tin-
side the young vascular bundles, which will later determine the venation, are
;

indicated. C, fully-grown leaf, with the course of the vascular bundles in-
dicated diagrammatically. D, a transverse section of the basal portion of a bud
showing three vascular bundles in each leaf. ./,', a similar section at a higher level ;

the number of vascular bundles has increased by branching. (After 1 (KIXKOA. from
GOEBEL'S Organography. A, li, and K slightly magnified.)

simple, as in the needles of Conifers, the primordial leaf simply

increases in length without any further differentiation into parts. In
other undivided leaves, however, whether lanceolate, elliptical, ovate,
or otherwise shaped, the flat leaf-blade is distinct from the leaf-base,
while a leaf-stalk may also be interpolated between them. If no
leaf-stalk developed the leaf is said to be SESSILE, otherwise it is
is

described as STALKED.
SECT. I MORPHOLOGY

Sessile leaves usually clasp the stem by a broad base. Where, as in the
case of the Poppy (Papaver somniferum), the leaf-base surrounds the stem, the
leaves are described as AMPLEXICAUL if, as in species of Bupleurum, it com-
;

pletely surrounds the stem, the term PEHFOT.I.VTK is used. If the bases of two
opposite leaves are united, as in
the Honeysuckle (Louicera Capri-
folium), they are said to be CON-
NATE. Where the blade of the
leaf continues downwards along the
stem, as in the winged stems of
the common Mullein (ferbaseum
fhapsiforme), the leaves are dis-
tinguished as DECURRENT. The
petiole of a leaf merges either
directly into the leaf-base, or it

swells at its lower end into a

LEAF -CUSHION Or PULVINUS, and
is thus articulated with the leaf-
base. This is the case, for in-
stance, with many of the Legu-
minosae. The leaf-blade, in turn,
may be either sharply marked oH'
from the petiole, or it may be pro-
longed so that the petiole appears
winged, or again it may expand
at its junction with the petiole into
ear-like lobes. A leaf is said to
be ENTIRE if the margin of the
leaf-blade is wholly free from in-

dentations, or if the latter are very

shallow. When the incisions are
deeper, but do not extend half-way
to the middle of the leaf-blade,
a leaf is distinguished as LOBKD ;

when they reach more than half-

way, as CLEFT (Fig. 35 sb) if the ;

incisions are still deeper the leaf

is said to be PARTITE (Fig. 38 I),
and they penetrate to the midrib
if

or base of the leaf- blade it is termed

DIVIDED. The divisions of the leaf- Kio. 34. Lily of th Valley (Convallariu mcjalis). ntl,
Scale leaves ; Ib, foliage leaves ; hb, bracts 6, flower ;
;

blade are said to be PINNATE or

rhizome aw, adventitious roots. (Somewhat re-
i'.-.-.-, ;

PALMATE, according as the incisions duced.)

run towards the midrib or towards
the base of the leaf-blade. Where the divisions of the leaf-blade are distinct and
have a separate insertion on the common leaf-stalk or on the midrib, then termed
the SPINDLE or RHACHIS, a leaf is spoken of as COMPOUND (Fig. 35 ub) in all ;

other cases it is said to be SIMPLE. The single, separate divisions of a com-

pound leaf are called leaflets. These leaflets, in turn, may be entire, or may be
divided and undergo the same segmentation as single leaves. In this way double
and triple compound leaves may be formed. Simply and doubly PINNATE leaves
in which the leaflets are attached to the two sides of the rhachis are of common
D
 BOTANY

occurrence. The leaflets are either sessile or stalkeil and sometimes also, as in
;

Knit!, tin and Mimosa, their stalks articulate with the spindle by means of swollen
pulvini. The term I-EDATK is ap-
plied to leaves in which the seg-
ments are further divided on one
side only, and the new segments
are similarly divided (Fig. 38 I).
Variations in the outline of leaves,
whether they are entire, serrate,
dentate, crenate, incised, etc., as
well as peculiarities in their ,-hapc
and segmentation, are of use in the
determination of plants.

A system of strands
known as the VEINS or

Flo. 35. Ranunculus aquatilis. ub, Submerged leavs ;

sb. floating leaves ; l>,

flower ; /, fruit. (Reduced.)

NERVES, traverses the flat leaf-blade. They

project more or less from the surface, especi-
ally on the lower side 'where they often
form definite ribs. Frequently the nerve
in the median line of the lamina is more

strongly developed and is then termed the

midrib ; in other cases several equally de-
veloped main nerves are present. Lateral
nerves spring from the one or more main
nerves.
l-'iu. ::i''. -Part of >tem and leaf <if

According to their VENATION or NEUVATUUE ;i .urass. //, Haulm: r, leaf-

leaves are described as PAUAU.KI, VEINED or NETTED sheath k, swelling of the Irut-
;

VEINED. In parallel venation the veins or nerves sheath above the node s, part ;

of leaf-blade /, ligule, (Nat.

;
run either approximately parallel with each other
sixe.)
or in curves, converging at tfie base and apex of the
leaf (Fig. 36 s) ; in netted veined leaves (Fig. 190) the veins branch olf from one
another, and gradually decrease in size until they form a fine anastomosing
network. In leaves with parallel venation the parallel main nerves are usually
SECT, i MORPHOLOGY 35

unitril liy weaker cross veins.

Netted or reticulately veined leaves in which tin-
side veins run from the median main nerve or MIDRIB are further distinguished
as I'iNNATKLY VEINED, or as PALMATELY VEINED when several equally strong ribs

separate at the base of the leaf-blade, and give rise in turn to a network of weaker
veins. Parallel venation is characteristic, in general, of the Monocotyledons ;

reticulate venation, of Dicotyledons. Monocotyledons have usually simple leaves,

while the leaves of Dicotyledons are often compound, and are also more frequently
provided with stalks.
The nerves or veins give to a leaf its necessary mechanical rigidity and render
possible its flattened form. The branches of the veins parallel to the margin of
most leaves prevent their tearing when there are no such marginal nerves in large
:

thin leaves, the lamina is easily torn into strips by the wind and rain. This fre-
quently happens to the leaves of the Banana (Musa), which, consequently, when
growing under natural conditions in the open air, presents quite a different appear-
ance from what it does when grown under glass. The leaves of the Banana, after
becoming thus divided, offer less resistance to the wind. In a similar manner the
leaves of Palms, although undivided in their bud state, become torn even during the
process of their unfolding. A similar protection from injury is afforded to the
Aroid (Monstera) by the holes with which its large leaf-blades become perforated.
Equally advantageous results are secured by many plants whose leaves are, from
their very inception, divided or dissected. The submerged leaves of aquatic
plants, on the other hand, are generally finely divided, not only for mechanical
purposes, but also to afford a more complete exposure of the leaf surface to the
water. Accordingly, in such water-plants as Ranunculus aquatilis (Fig. 35), which
possess both floating and submerged leaves, it is generally the latter only that are
dissected and The pointed extremities (DRII> TIPS) of the
filiform in character.

foliage leaves of land plants, according to STAHL ( 18 ), facilitate the removal

many
of water from the leaf surface. Fleshy so-called succulent leaves, like fleshy stems,
serve as reservoirs for storing water.

Heterophylly. Many plants are characterised by the develop-

ment of different forms of foliage leaves. Such a condition is
known as heterophylly. Thus the earlier leaves of Eucalyptus
globulus are sessile and oval, while those subsequently formed are
stalked and sickle -shaped. In other cases the heterophyllous
character of the leaves may represent an adaptation to the surround-
ing environment, as in the Water Crowfoot (Ranunculus aquatilis), in
which the floating leaves are lobed, while those entirely submerged
are finely divided (Fig. 35). A
number of epiphytic Ferns belonging
to the genera Polypodium and Platycerium alternately bear stalked,
pinnate leaves serving for assimilation, and broad, sessile, nsually
cordate NEST-LEAVES which serve to collect humus ; when, as MANTLE-
LEAVES, they are closely appressed to the substratum they form
humus by their rapid decay. The humus thus collected or produced
19
is
penetrated by the roots of the Fern ( ). .

The Leaf -base. In Monocotyledons the leaf-base very often

forms a SHEATH about the stem in Dicotyledons this happens much
;

less frequently. In the case of the Gramineae, the sheath is open on

the side of the stem opposite the leaf-blade (Fig. 36 r), while in the
36 BOTANY PART I

Cyperaceae it is completely grown together. The sheath of the

Grasses is prolonged at the base of the lamina into a scaly out-
growth, the ligule (/). Such a sheath, while protecting the lower
part of the internodes, which remain soft and in a state of growth,
gives them at the same time rigidity. STIPULES are lateral
appendages sometimes found at the base of leaves. When present
they may be either small and inconspicuous (Fig. 37 nb), or may
attain a considerable size. When their function is merely to
protect the young growth in the bud, they are usually of a brown
or yellow colour, and are not persistent whereas, if destined to;

become assimilatory organs, and to assist in providing nourishment,

FIG. 37. ttin\Cherry(l'runusAviuin). FIG. 3S.Helleboru.- fott'idu*. Foliage leaf

Bud-scales (1-3) and the transition (Oiaiul intermediate forms U-tweeii this
forms (4-6) to the foliage leaf (7),; and the bract (h). (Reduced.)
sp, leaf -blade; s, leaf-stalk; nb,
stipules. (Reduced slightly.)

they are green, and may assume the structure and form of the
leaf-blade, which sometimes becomes modified and adapted to other
48, 49). Normally, the stipules are two in number,
purposes (Figs.
that is, one on each side of the petiole. In many species of Galium,
where*the stipules resemble leaf-blades, the leaf-whorls appear to be
two leaves
composed of six members, but consist actually of but
with their four stipules, which may be easily distinguished by the
absence of buds in their axils. In other species of the same
any
are only four members in
genus (Galium cruciatum and palujtre) there
the whorls, as each two adjoining stipules become united. In many
cases, as in the Rose and the Clover, the stipules
have the form of
appendages to the enlarged leaf-base. Sometimes
both stipules are
united into a single one, which then appears to have an axillary
 MORPHOLOGY 37

origin or the stipules may completely encircle the stem, and thus
;

form a sheath (OCHREA) about the younger undeveloped leaves.

This sheath-like fusion of the stipules may be easily observed on the
India-rubber tree (Ficus elastica), now so commonly grown as a
decorative plant. In this case the stipular sheath is burst by
the unfolding of each new leaf and pushed upwards. In the
Polygonaceae the stipular covering is similarly torn apart by the
developing leaves, but remains on the stem in the form of a
membranous sheath.
Scale Leaves possess a simpler form than foliage leaves, and are
attached directly to the stem, without a leaf-stalk. They exercise
no assimilatory functions, and are more especially of service as
organs of protection. Scale leaves exercise their most important
function as BUD-SCALES (Fig. 37); they are then hard and thick, and
usually of a brown colour. They most frequently take their origin
from the enlarged leaf-base in that case the upper leaf either does
;

not develop, or exists only in a reduced condition at the apex of

the scale. The true morphological value of scale leaves of this
nature is very evident in the bud-scales of the winter buds of the
Horse-chestnut (Aesculus Hippocastanum) for, while the outer scales
;

show no perceptible indications of an upper leaf, small leaf-blades

can be distinctly distinguished at the apices of the inner scales. In
other cases the scale leaves are modified stipules (Fig. 37), and are
then also derived from the leaf-base ; while, in other instances, they
correspond to the enlarged, but still undifferentiated, primordial leaves.
The bud-scales of the Oak are the stipules of leaves of which the
laminae are only represented by minute scales. Scale leaves,
usually colourless and in various stages of reduction, are found on
rhizomes (Figs. 23, 34), bulbs (Fig. 25), and tubers (Fig. 26). On
the aerial stems arising from such subterranean shoots the forma-
tion of similar scale leaves generally., precedes the development of
1

the foliage leaves, with which they are connected by a series of

transitional forms.
Braeteal Leaves resemble scale leaves in form, and have a
similar origin (Fig. 34 hb). They act as subtending leaves for the
floral shoots, and are termed BRACTS. They are connected with
foliage leaves by intermediate forms (Fig. 38). Though they are
not infrequently green they may be otherwise coloured, or even
colourless.
Floral Leaves. The modified leaves which form the flowers of
Phanerogams are termed floral leaves. In the highest development
attained by a phanerogamic flower (Fig. 39), the successive floral
leaves are distinguished as sepals (&), petals (c), stamens (), and
carpels (<?). In form the sepals and petals approach the bracts.
In most cases the sepals are green and of a firm structure the ;

petals, on the other hand, are more delicate and variously coloured.
 BOTANY

The stamens are generally filamentous in shape, and produce the

pollen in special receptacles. The carpels more closely resemble
scale leaves, and in Angio-

sperms by closing together

form receptacles within which
the ovules are produced.
The stamens and carpels
of Phanerogams correspond
to the spore-bearing leaves
of the Vascular Cryptogams.
Such -
spore bearing leaves
are termed SPOROPHYLLS,
and even in ,the Vascular
-Jc
Cryptogams exhibit a greater
or less departure from the
form of other foliage le;i\c>.
Kiu. Klower of Pneoniii />r/e/rtrta.
:!'.i.
Sepals; c,
^
/.',

petals stamens a, carpels. Some of the sepals,

; ;

petals, ami stamens have been removed to show the

k {s cyi(lent fhafc ^ 1(
.

lacteal leaves are to be considered

pistil, consisting of two serrate carpels. (Half nat.
size.) as rudimentary foliage leaves, not

only from the mode of their de-

velopment but also from the possibility of transforming them into foliage leaves.
GOEBKL, by removing the growing tip and foliage leaves of a shoot, succeeded in
20
forcing it to develop other foliage leaves from its scale leaves ( ). Rhizomes,
grown in the light, develop foliage leaves in place of the usual scale leaves, and
even on a potato it is possible to induce the formation of small foliage leaves
instead of the customary scale leaves.

Leaf -Sears. After a leaf has fallen, its previous point of

insertion on the stem is marked by the cicatrix or scar left by the
fallen leaf.In winter, accordingly, when the trees are denuded of their
leaves, the axillarybuds are seen to be situated above the leaf-scars.
Vernation and ^Estivation. * A section through a winter bud
shows a wonderful adaptation of the young leaves to the
narrow space in which they are confined. They may be so disposed
that the separate leaves are spread out flat, but more frequently
they are folded, rolled (Fig. 40 /), or crumpled. The manner in
which each separate leaf is disposed in the bud is termed VERNATION*.
On the other hand, the arrangement of the leaves in the bud with
respect to one another is designated .-ESTIVATION. In this respect
the leaves are distinguished as FREE when they do not touch, or
VALVATE when merely touching, or IMBRICATED, in which case some
of the leaves are overlapped by others (Fig. 40 /). If, as frequently
occurs in flower-buds, the margins of the floral leaves successively
*
[The use of these terms in the following paragraph differs from that customary in
England. By VERNATION is understood the arrangement of the leaves in a vegetal i\<-
hud as a whole. The folding of each individual leaf in the Imd is termed ITYXIS. The
term .ESTIVATION is applied to the arrangement of the parts in a Hower-lmd. TRANS.]
 MORPHOLOGY

overlap each other in one direction, the aestivation is said to be

CONTORTED.
The Arrangement of Leaves. In all erect elongated shoots, and
still more so in dwarf shoots, it is apparent that there is a marked

regularity in the arrangement of leaves. This regularity may be

most easily recognised in cross-sections of buds (Fig. 41), particularly
in sections showing the apex of the vegetative cone (Figs. 31, 33),
From such evident that the developing leaf-rudiments
sections it is

stand in the relative position to the pre-existing leaves which

21
best utilises the available space. According to Schwendener ( )
the arrangement of the leaves on the axis is determined by purely
mechanical causes it is dependent on the pressures and tensions
;

FIG. 40. Transverse section of a bud of Fio. 41. Transverse section of a leaf-bud
a lit ni.
k, Bml-scales showing imbricated of Tsugii<<<
iiiidensis, just below the
ifstivation [vernation] ; I, foliage leaves with apex of the shoot, showing a -f3 diver-
involute vernation fptyxis] a, each leaf has
; gence. (After HoKMKISTER.)
two stipules, (x 15.)

induced by the leaves which always stand in contact at their origin.

The original arrangement of the young leaves may be modified as
growth continues owing to the mutual pressure they exert. ) If
the axis does not grow in length, but only in thickness, as the
rudimentary leaves increase in size, their points of insertion will be
displaced laterally by longitudinal pressure ; if the axis increases in
length, and not in thickness, the insertion of the leaves will be
displaced by a transverse pressure. The arrangement of the leaves
would also be affected
by any increase or decrease in the size of the
vegetative unaccompanied by a corresponding increase or
cone,
cessation of the growth of the rudimentary leaves.
In developing flowers sudden changes in the relative position of
the parts often occur in which the apical cone undergoes rapid
increase in size while the leaf rudiments become smaller. Changes
in the usual position of the leaves may also be occasioned by the
40 BOTANY

torsion of their parent Thus the leaves of Pandanus first

stein.

appear in three
straight rows on the vegetative cone, and their
subsequent spiral arrangement results from the torsion of the stem.
Irregular arrangement of the leaves, such as occurs, for example, on
the flower-stalk of the Crown Imperial (Fritillaria imperialis), may
result from the unequal size of the leaves at the time of their
inception on the vegetative cone. Further observation has revealed
an increasing number of cases in which definite leaf arrangements
cannot be explained by the spatial relations, and as the effect of
contact and pressure. It can only for the present be assumed that
the cause of the leaf-arrangement in these cases lies in the inherited
organisation of the plant ().
A
frequent mode of arrangement of foliage leaves is the decussate,
in which two-leaved whorls alternate with each other (Figs. 31, 33).
A whorled arrangement is characteristic of floral
leaves. When the number
of leaves in each
whorl the same the whorls usually alternate.
is

On the other hand, the number of members in

the different whorls of floral leaves will often
be found to change, especially on passing from
the petals to the stamens, or from the latter to
the carpels. In many cases a whorl, the existence
of which would be expected from the position of
other whorls and from a comparison with allied
Fl - Dia8ram r a
plants, may * be altogether wanting. In this con-
:
Liliaceous (lower. The r
.,; '
. * i T
main axis is indicated nection a comparison of the flowers of the Liliaceae
by a black dot, oppo- an(j Iridaceae will be instructive. The flowers of
site to which is the .1 T .
/& . \ j n
bract
the Liliaceae (.tig. 42) are composed of five
regularly alternating, three-leaved whorls or cycles,
viz. a calyx and a corolla (each consisting of three leaves, and, on
account of their similar appearance, usually referred to conjointly as
the PERIANTH), an outer and an inner whorl of stamens, and finally,
in the centre of the flower, an ovary of three carpels. In the flowers
of the Iridaceae (Fig. 43) the arrangement is exactly similar, except
that one whorl, that of the inner cycle of stamens, is lacking, but
the three carpels are situated exactly as if the missing whorl of
stamens were present. From this similarity of arrangement,
despite the absence of the one cycle of stamens, the conclusion
has been drawn that, at one time, the inner row of stamens
was actually present, but has now disappeared. In constructing
a THEORETICAL DIAGRAM of the Iridaceae the missing cycle of
stamens is
by some special sign (by crosses in Fig. 43)
indicated ;

a diagram which theoretical suppositions are not taken into

in
consideration is called an EMPIRICAL DIAGRAM. Diagrams showing
the alternate arrangement of leaves, in cases where only a single
leaf arises from each node, may be constructed by projecting the
 MORPHOLOGY 41

successive nodes of a stem upon a plane by means of a series of

concentric which the position of the leaves may be
circles, on
indicated (Fig. 44). The angle made by the intersection of the
median planes of any two successive leaves is called
their DIVERGENCE, and is expressed in fractions of
the circumference for example, when the angular
;

divergence between two successive leaves is 120,

their divergence is expressed by the fraction |.
In the adjoining diagram (Fig. 44) a f divergence
is shown. Where the lateral distance between
two successive leaves is of the circumference of
-?-

the stem, the sixth leaf is above the first, the

seventh above the second, and so on. The leaves FIG. 43. Theoretical
form on the axis five vertical rows, which are diagram of the flower
Where the leaves of the Iris. The ab-
spoken of as ORTHOSTICHIES. sent whorl of stamens
are very much crowded, as in dwarf-shoots, a is indicated by
set spiral rows called PARASTICHIES, due to
of crosses.

the contact of the nearest laterally adjacent mem-

bers, becomes much more noticeable than the orthostichies. If the
surface of such an axis be regarded as spread outl horizontally,
the parastichies become at once distinguishable (Fig. 45), and it will
be evident that the sum of the parastichies cut by every cross-section

FIG. 44. I
)iagram showing j position of Fio. 45. The \ position on the outspread
leaves. The leaves numbered according surface of the axis, o, Orthostichies p, ;

to their genetic sequence. parastichies. The leaves are numbered

according to their genetic sequence.

through such an axis must equal the number

of the orthostichies.
On objects like pine cones, in which the parastichies are easily
be used to determine the leaf arrangement.
recognised, they may
If a line be drawn on the surface of a stem, so as to pass in the
42 BOTANY PART i

shortest way successively through the points of insertion of every

leaf,a spiral called the GENETIC SPIRAL will be constructed. That
portion of the genetic spiral between any two leaves directly over
each other on the same orthostichy is termed a CYCLE. AVhere the
divergence is jj, a cycle will accordingly include five leaves, and will
in such a case have made two turns about the stem. The most
common divergences are the following, -\, V, i;,.',,
s
n ;>, etc.
,"'..,
., .'.

In this series it will be observed that in each fraction the numerator

and denominator are the sum of those of the two preceding fractions.
The value of the different fractions varies accordingly between ?,

and 7j, while always approaching a divergence angle of 137 30' 28".
The great majority of leaf arrangements can be expressed by the
terms of this main series of divergences.

This main series was discovered by CARL SCHIMPEK and ALEXANDER I.KAI v
It exhibits a rational relation of the divergences to the circumference of the axis,
so that, as the number of leaves increases, definite leaves are situated accurately
above one another. As WIKSXKU 2:1 ) in particular
(
has made clear, it ditl'ns
from all other possible series in attaining the most equal distribution of the
leaves on the axis bearing them, while requiring the smallest number of
leaves. This results in an advantageous utilisation of the available space, a well-
distributed loading of the axis and, when the latter is vertical, in the best utilisa-
tion of the illumination. The importance of these advantages as determining the
leaf-arrangement is seen in those cases in which a plant bears only a few (2-4)
leaves. These stand in a whorl at equal distances from one another and thus
their weight is equally distributed, and they obtain equal amounts of light.
When leaves are arranged alternately on a vertical axis, their size and shape,
together with the length of the iuternodes, ensure each obtaining the requisite
amount of light. This arrangement is not a convenient one, and as the leaves borne
on a vertical axis increase in number their divergence becomes progressively higher.
It is otherwise in inclined or horizontal axes here the divergence is relatively
;

low, usually or an approximationto this, since this corresponds to the most

favourable exposure of the leaves to the light. In most instances this advan-
tageous result is attained by_twisting of the internodes thus when, as is often
;

the case, the leaves are decussate on an erect axis, they form four vertical rows ;

but when it is inclined they are brought by twisting to stand in two rows.
Similar secondary changes exposing the leaf-blades to the light affect alternately
arranged leaves. The position of the foliage leaves is indeed always clearly
adapted to the need of illumination. When the leaves form a rosette, the stalks
of those lowest on the stem are frequently elongated, so that their blades are not
shaded by the more central leaves. This is especially well seen in the floating
rosettes of Trapa natans.
While the arrangement of the foliage leaves conforms on the whole to the main
series of divergences, this is not usually suitable in the case of the foliar structures
of flowers and inflorescences, which have different purposes to serve. Other
relations of position also occur in the vegetative region, as has been shown l>y
2
GOEBEL C *), in particular in the case of dorsiventral shoots. The tips of
dorsiventral shoots are frequently coiled ventrally inwards, bearing their leaves
either dorsally or on the sides, but, in the latter case, approaching the dorsal
surface. The creeping stems of many Ferns or the flower-bearing shoots of
 MORPHOLOGY 43

Forget-me-not (ifyvstts) are good examples of such dorsiventral shoot.-. Tlie lint-

joining successive leaves in such case is, at the best, but a zigzag. On the
ground of such observations as these it may be concluded that the actual leaf-
arrangements represent adaptations to definite conditions of life, and that with
alterations in the latter other arrangements must arise.

The Metamorphosis of Foliage Leaves. A striking modification

of foliage leaves is seen in peltate leaves, in which the petioles are
attached to the lower surfaces somewhat within the margin, as in
the leaves of the Indian
Cress (Tropaeolum majus,
Fig. 191). In the process
of their development
the young leaf -blades, in
this case, grow not only
in the same direction as
the petioles, as a prolonga-
tion of them, but also hori-
zontally in front of them.
The tubular leaves of many
insectivorous plants may
have commenced their de-
velopment in much the
same way. The leaves of
'Nepenthes robusta (Fig. 46),
for example, in the course
of adaptation to the per-
formance of their special
function, have acquired the
form of a pitcher with a
lid which is closed in
young leaves, but eventu-
The pitcher, as
ally opens.
GOEBEL has shown, arises
as a modification of the
leaf-blade. At the same
time the leaf-base becomes
expanded into a leaf-like I-'K;. 4ii. Xi-penthes robuxtc. iial. sixr.)
(.',

body, while the petiole

between the two parts sometimes fulfils the office of a tendril.
By a similar metamorphosis of its leaflets, bladder-like cavities
are developed on the submerged leaves of Utricularia (Fig. 47).
The entrance to each bladder is fitted with a small valve which
permits the ingress but not the egress of small water-animals.
While such leaves display a progressive metamorphosis the
modification may be of the nature of a reduction, as is the case
in many Ferns, which form leaf-runners. Like the modified
44 BOTANY

shoots of the same name these are elongated and enable the
bud produced at the end to develop at a distance from the parent
plant. Camptosorus rhizophyllns, an American Fern, is commonly
known as the Walking Fern. Such leaf-runners usually lose their
pinnae and are reduced to the leaf stalk. A particularly striking
appearance is presented in those cases in which the first leaf of the
bud in its turn forms a leaf-runner, so that a sympodium of runners
results (Asplenium obtusifolium, A. Mannii).
Ametamorphosis of the whole leaf lamina, or a part of it, into
tendrils (LEAF-TENDRILS) is of comparatively frequent occurrence,

Kic. 17. l'tricu!<iriii~>-u.l<jari*. A, 1'art of leaf with several bladders (x 2). 11, Single pinnule of
leaf with bladder (x 6). C (after GOEBEL), Longitudinal section of a bladder (x 28)
; .valve;

", wall of bladder.

especially among the Papilionaceae. In the leaf of the Pea (Fig. 48),
the leaflets of the upper pairs have become transformed into delicate
tendrils which have the power of twining about a support. In the
case of the yellow Vetchling, Lathyrus Apliaca (Fig. 49), the whole
leaf is reduced to a tendril and the function of the leaf -blade is
assumed by the stipules (ri). A
comparison between these two forms
is instructive, as it indicates the steps of the gradu-
phylogenetically
ally modifying processes which have resulted in the complete reduction
of the leaf lamina of Lathyrus. The comparison of the two preced-
ing cases with Ampelopsis (Fig. 29) will make the distinction between
these tendrils and stem-tendrils clear, and indicate the value of
comparative morphological investigation.
In Lathyrus Aphaca the stipules assume the function of the meta-
 MORPHOLOGY 45

morphosed leaf laminae ; in other instances, as in the case of the

Australian species of Acacia (Fig. 171, 7, 8, 9), it is the leaf petioles
which, becoming flattened and leaf-like in appearance, supply the place
of the undeveloped leaf-blades. Such a metamorphosed petiole is called

FIG. 48. Portion of stem and leaf of the common Pua, Fie. i'.'. l.at/ii/nt-s .I
J'isum sativuin. *, Stem,; n, istipules ; b, leaflets of *, Stem ; n, stipules;

the compound leaf; r, leaflets modified as tendrils ; , b, leaf-tendril. ( nat.

floral shoot, (i nat. size.) size.)

a PHYLLODE, and, except that it is expanded perpendicularly, closely

resembles a cladode. From the latter, however, it is morphologically
different, for the one represents a metamorphosed petiole, the other a
metamorphosed shoot. In accordance with
this distinction phyllodes do riot, like cladodes,

spring from the axils of leaves. Just as

stems become modified into thorns (Fig. 30),
by a similar metamorphosis leaves may be
converted into LEAF THORNS. Whole leaves
on the main axis of the Barberry (Berberis
vulgaris) become thus transformed into thorns,
usually three, but in their character of leaves
still subtend axillary shoots provided with

foliage leaves. By a similar metamorphosis,

the two stipules of the leaves of the common
Locust (Eobinia Psewlacacia) become modified
into thorns, while the leaf lamina persists
FIG. 50. Part of stem and com-
as a foliage leaf (Fig. 50).
pound leaf of Sobinin Pseud-
ncacin. n, Stipules modified
The Root into thorns ;/, pulvinus.
:

(A nat. sixc.)
The third primary member of the corruo-
as an underground root
phytic plant in its typical development
less marked differences in external form than were shown by
presents
46 BOTANY i-.uir i

the other members. This may be put in relation with the uniform
conditions to which roots are exposed in the soil. Certain differences
are, however, found between the roots of plants adapted to live in
25
different situations ( ). The root has as its most important function t la-
absorption of water and nutrient substances dissolved in it from tin-
soil, and also serves to attach the plant firmly in the soil. Frequently it
serves for storage of assimilated substances. The general appearance
of an underground root differs as it is more specially adapted to perform
one or other of these functions. Those roots which grow in water or
mud become more elongated, are little branched, and are often pro-
vided with special arrangements for aeration which lead to localised
swellings ; they do not, however, undergo any fundamental change
of form. Those roots which grow in the air, AERIAL ROOTS, tend t<>
be more strikingly modified.
The absence of leaves and the existence of a root-cap protecting
the growing point are characteristic of roots, and furnish an easy
means of distinguishing them from underground shoots. A ROOT-CAP or
CALYPTRA affords the vegetative cone of a root the protection that is
provided to the apex of a stem by the leaves of the bud. Although,
generally, the existence of a root-cap is only disclosed by a median,
longitudinal section through the root-tip, in some roots it is plainly
distinguishable as a cap-like covering. The very noticeable cap- mi
the water roots of Duckweed (LemtM) are not, in reality, root-caps,
as they are not derived from the root, but from a sheath which
envelops the rudimentary root at the time of its origin. They are
accordingly termed ROOT-POCKETS. As a general rule, however, roots
without root-caps are of rare occurrence, and in the case of the
Duckweed the root-pockets perform all the functions of a root-cap.
The short-lived root of the Dodder (p. 27) affords another example
of a root devoid of a root-cap. Characteristic of roots are also the
ROOT-HAIRS (Fig. 170, r), which are found at a short distance from
their tips. As the older root-hairs die at the same rate that the
new ones are developed, only a small portion of a root is provided
with root-hairs at the same time. In some few instances roots
develop no root-hairs; this is true of the roots of many Conifers.
Branching of the Root. Just as the shoot may become bifurcated
by the division of its growing point (Fig. 18), so a root may become
similarly branched. For the most part, this mode of branching
takes place only in the roots of Lycopodiaceae, the shoots of which are
also dichotomously branched (p. 19). The branches of roots usually
occur in acropetal succession, but the lateral roots (Fig. 170 w) make
their appearance at a much greater distance from the growing point
of the main root than lateral shoots from the apex of their parent
stem. By reason of the internal structure of their parent root, lateral
roots always develop in longitudinal rows (Fig. 1
70). They are of endo-
genous origin, and before reaching the surface must break through the
 .MORPHOLOGY 47

surrounding and overlying tissue of the parent root, by the ruptured

portions of which they are often invested at the base, as with a collar.
ADVENTITIOUS HOOTS, just as adventitious shoots, may arise from
any part of a plant. They are especially numerous on the under side
of rhizomes (Fig. 23 w), and also, when the external conditions are
at all favourable, they seem to develop very readily from the stem
nodes. A young shoot, or a cutting planted in moist soil, quickly
forms adventitious roots, and
roots may also arise in a similar
manner from the bases of
leaves, especially from Begonia
leaves when planted in soil.
The origin of adventitious roots,
as of almost all roots, is en-
dogenous. Dormant root rudi-
ments occur in the same manner
as dormant buds of shoots.
Willow-twigs afford a special
case of the presence of such dor-
mant rudiments of adventitious
roots, the further development
of which is
easily induced by
darkness and moisture.
When adventitious shoots
are borne upon roots they
Fin. 51. Root-tubers of iJahlia rariabilis. The
mostly arise at some distance lower portions of tin- cut stems. (! nat.
from the apex. In the Adder's
Tongue Feru (Ophioglossum), the vegetative reproduction of which is

by means of buds formed on the root, these arise

effected close to the

root-apex. Lastly, just as terminal leaves may arise from the apex of
a stem and terminal shoots from the tips of the leaves of Ferns, in a
few instances (species of Asplenium and Platycerium) a shoot may arise
as the direct continuation of the growing point of a root ( 2r> ).
The Form of Subterranean Roots. The customary nomenclature
for the various root based on their shape, size, and mode of
forms is

branching. A a prolongation downwards of the main

root which is

stem is called the main root or TAP-ROOT the other roots are termed,
;

with reference to the tap-root, LATERAL ROOTS of different orders,

according to the order of their development. A tap-root is present in
Gymnosperms and many Dicotyledons, while it is,as a rule, wanting
in Monocotyledons. The root system
most Monocotyledons and
of

many Dicotyledons mainly from

consists of adventitious roots arising
the base of the stem or from underground stems or rhizomes. Roots
which serve for storage of reserve materials may enlarge and become
turnip-shaped or tuberous (Fig. 51). Such tuberous growths often
greatly resemble stem tubers, but may be distinguished from them
48 BOTANY

by their root-caps, by the absence of any indications of leaf develop-

ment, and by their internal structure.

The morphology of the tubers of the Orchidaceae is peculiar. They are, to

a great extent, made up of flesh}-, swollen roots, fused together and terminating
above in a shoot-bud. At their lower extremity the tubers are either simple or
palmately segmented. In the adjoining ligure (Fig. 52) both an old (f) and a
young tuber are represented still united together.
(t") The older tuber has pro-
duced its flowering shoot (b), and has begun to shrivel a bud, formed at the base
;

of the shoot, in the axil of a scale leaf (s), has already developed the adventitious
roots, which, swollen and fused together, have given rise to the younger tuber.
Roots of ordinary form arise from the base of the stem above the tuber.

Metamorphosis of Roots. The aerial roots of tropical EPI-

PHYTES C27 ) differ considerably in their structure from underground
roots. The aerial roots of the Orchidaceae and of many Aroideae are
provided with a spongy sheath, the VELAMEN, by means of which
they are enabled to absorb moisture from
the atmosphere. Aerial roots, in some cases,
I
grow straight downwards, and upon reaching
the ground, branch and function as nutritive
roots for the absorption of nourishment ; in
other instances, they turn from the light,
and, remaining comparatively short and un-
branched, fasten themselves as CLIMIJINC;
ROOTS to any support with which they come
in contact. The climbing roots of many
Orchids, Aroids, and Ferns branch and form
r The
lodgment *places for humus: and into this
Fio.52.-o,rM*taft/w,-,,.
old root-tuber; t", the young , , , , , '. ,.
root-tuber; b, floral
shoot; s,
absorbent branches of the climbing roots
scale leaf with axillary bud,
penetrate. Pendent aerial roots generally
A-, from which the new tuber
contain chlorophyll in their cortical tissue.
has arisen ; r, ordinary adven-
titious roots, (i nat. si/.'.) InOrchids belonging to the genera, Angraecuin
and Taeniophyllum, the task of nourishing the
plant is left entirely to the aerial roots, which are then devoid of a
velamen, and very much flattened. They are distinctly green-
coloured, and supply the place of the leaves which lose their green
colour and are reduced to scales. The flat, dorsiventral, chlorophyll-
28
containing roots of the tropical Podostemaceae ( ), which have been
referred to with regard to the sequence of shoots (p. 29), fulfil a
similar function. The aerial roots of the epiphytic Bromeliaceae
are developed exclusively as climbing roots, while the leaves function
not only as assimilating organs, but also assume the whole task of
water-absorption. All the aerial roots of epiphytes are, so far as
their originis concerned, adventitious.

The numerous adventitious roots which form a thickly matted covering on the
trunks of Tree-ferns become hard after death, and serve as organs of protection.
 MORPHOLOGY 49

In some Palms (Acantkarrhiza, Iriartca) the adventitious roots on the lower part
of the stem become modified into thorns, ROOT-THORNS. The roots of certain
tropical plants, such as Paiidanus and the swamp-inhabiting Mangrove trees, are
specially modified. These plants develop on their stems adventitious roots, which
grow obliquely downwards into the ground, so that the stems finally appear as if
standing on stilts. The Banyan trees of India (Ficus indica) produce wonderful
root-supports from the under side of their branches, upon which they rest as upon
columns. The lateral roots of certain Mangrove trees become modified as peculiar
breathing organs, and for this purpose grow upwards into the air out of the
swampy soil or water in which the trees grow, and are provided with special
aerating passages. Such KESPIKATORY or AERATING ROOTS surround the Mangrove
trees looking like vigorous Asparagus stalks, and enable the roots growing below in
the mud to carry on the necessary exchange of gases with the atmosphere. (Cf.
Fig. 213.)

Reduction of Roots. There is "a general relation between the

degree of development of the leaf-surface and of the root-system.
In saprophytic and parasitic plants, the shoots of which are as a rule
extremely reduced, a corresponding reduction of the root-system can
be recognised. Special absorbent organs or HAUSTORIA are often
developed on the roots of parasites, e.g. on the roots of the green
partial parasites Euphrasia, Odontites, Thesium, etc., which become
attached to the roots of their host plants by means of disc-shaped
or wart-shaped haustoria. In extreme cases no subterranean roots
may be developed, and the parasite only bears haustoria which
penetrate the host (Cuscuta, Fig. 202 H). The haustoria of the
Kafflesiaceae traverse the body of the host -plant as elongated
filaments of cells, and are capable of regenerating the parasite.
The immense flowers of Rafflesia Arnoldi, which spring directly from
the roots of Cissus, originate from similar haustoria. The reduction
of the roots may extend to such a degree that, in a number of plants,
no roots are formed. It has been already mentioned that in the
case of Coralliorrhiza innata (Fig. 24) the rhizome assumes all the
functions of the roots, which are entirely absent. Also in many aquatic
plants (Scdvinia, Wolffia arrhiza, Utricularia, Ceratophyllum) roots are
altogether absent since these plants no longer require them. The
beard-like epiphytic Tillandsia usneoides, belonging to the Bromeliaceae,
has no roots and obtains the water it requires by means of special
scale-like hairs.

Members of Independent Origin

Theexistence of parts of the plant with the characters of
distinctprimary members, but to which an independent phylognetic
29
origin must be ascribed, has been pointed out by GOEBEL ( ). The
tubers which occur among the Dioscoreaceae, an order of twining
plants belonging to the Monocotyledons, are of this nature (Fig. 53).
The extreme forms of these tubers are flat, cake-shaped bodies, which
in development and structure exhibit a mixture of the characteristics
E
50 BOTANY

of stem and root. The rhizophores of Selaginella (Fig. 54) which

have been variously interpreted as leafless shoots, as roots, and as
organs sui generis, also find their place here. They resemble leafless
shoots, attain a considerable length, and may branch dichotomously
and give rise to endogenous roots close to the growing points. As a
matter of fact they are in structure and general behaviour inter-
mediate between shoot and root, and it is not probable that they
have originated from either of these primary members.

FIG. 53. IKoscorea sinuata. Tuber separated from the shoot A, from above
;
: /;. from below.
The lower side bears roots.(After K. GOEBEL. A nat. sixc.)

Various outgrowths of the body of the plant to which an inde-

pendent origin is to be ascribed are grouped together as EMERGENCES.
A phylogenetic significance cannot be attached to the term as here
used. It includes structures of the most different origin belonging
both to the thallus and the cormus. The rhizoids, which serve to
fasten the thallus of many Thallophytes and the cormus of the
cormophytic Bryophyta to the substratum, as well as the massive
attaching organs of many Brown Sea-weeds (Fucaceae and Lami-
nariaceae) are classed here. So also are the structures which contain
the asexual and sexual reproductive cells of the cryptogams (sporangia
 MORPHOLOGY 51

and sexual organs). In the sense of the term implied here the hairs,
prickles, and glands borne on the surface of the highly organised
plants must be included as well as the haustoria described above.
As an extreme case the attaching organs (hapterae) of the previously
mentioned Podostemaceae may be referred to. These hapterse serve

Fio. 54. Part of the shoot of Sflaginella, Martensii with rhixo]>li< >*.

(From GOEBEL, Organography. Nat. size.)

to attach the nutritive roots of these plants firmly to the rocks

exposed to rapidly flowing water, upon which they grow They are
at first conical outgrowths, but flatten out and become lobed when
applied to the surface of the rock. The shoots originate from the
flat nutritive roots. There is, indeed, no reason why outgrowths of
the vegetative body of the plant should not become adapted to the
30
performance of particular functions ( ).

^-v
LIBRARY
52 BOTANY

II. INTERNAL MORPHOLOGY

(Histology and Anatomy

A. The Cell

1. STRUCTURE OF THK CKI.L

All plants and animals are composed of elementary organs

called cells. In contrast to animal cells, typical vegetable cells are
surrounded by firm walls, and are thus sharply
marked off from one another. In fact, it was
due to the investigation of the cell walls that
the cell was first recognised in plants. An
English micrographer, ROBERT HOOKE, was thr
first to notice vegetable cells. He gave them
this name in his Micrographia in the year 1667,
because of their resemblance to the cells of a
honeycomb, and published an illustration of a
piece of bottle-cork having the appearance shown
Km. 66. Copy of a part of /T ,. __ x
HOOKES illustration of m
. ,, . .

the adjoining figure (Fig. 55).

.

ROBERT -,-,

bottle-cork, which he dp- HOOKE, however, Was Only desirOUS of exhibit-

scribed as "Schematism or
texture of cork. o b meang f c]i fferent objects the
. L i
capabilities
i T
of his microscope ; consequently, the Italian,
MARCELLO MALPIGHI, and the Englishman, NEHEMIAH GREW, whose
works appeared almost simultaneously a few years after HOOKE'S
Micrographia, have been regarded as the founders of vegetable
histology. The living contents of the
cell,the real body or substance, was not
recognised in its full significance until
the middle of last century. Only then
was attention turned more earnestly to
this study, which has since been so especi-

ally advanced by MEYEN, SCHLEIDEN,

HUGO v. MOHL, NAGELI, DE BARY, FER-
DINAND COHN, PRINGSHEIM, and MAX
SCHULTZE. KK.. -,i;.-Knibryoiiir c.-ll from th.-

If an examination be made of a thin vegetative <! of ;,

longitudinal section of the apex of a stem

of a phanerogamic plant, with a higher cytoplasm-. -;,, t-hri)iuatoi>jior-s ;

'"' cel1 wal1 ( s< " >w '' ! " -I'^rain-

& power than that used in the
- l ''

magnifying
J. .
,. ,. ,,-,. 1fr . , matic, X circa 1000.)
previous investigation (rig. 17) of the
vegetative cone, it will be seen that it consists of nearly rectangular
cells (Fig. 56), which are full of protoplasm and
separated from one
another by delicate walls. In each of the cells there will be clearly
distinguishable a round body^&), which fills up the greater part of
 MORPHOLOGY 53

the cell cavity. This body is the cell NUCLEUS. If sections, made
in different directions through the vegetative cone, be compared with
one another, it will be seen that its component cells are nearly
cubical or tabular, while the nuclei are more or less spherical or
disc-shaped. The finely granular substance (cy) filling in the space
between the nucleus (k) and the cell wall (m) is the CELL PLASM or
CYTOPLASM. In the cytoplasm there are to be found, about the
nucleus, a number of colourless and highly refractive bodies these :

are the pigment-bearers or CHROMATOPHORES (ch). NUCLEUS, CYTO-

PLASM, AND CHROMATOPHORES, CONSTITUTE THE ELEMENTS OF THE
LIVING BODY OF A TYPICAL VEGETABLE CELL. To designate all
these collectively, it is customary to use the term PROTOPLASM,

which isthen to be understood as including all the living con-

stituents of the cell or PROTOPLAST.
In many animal cells modern investigations have revealed, in
addition to the constituents of the protoplasm just mentioned, a
small structure situated close to the
nucleus, which has been termed the CEN-
TROSOME, CENTRIOLE, or ATTRACTION-
SPHERE. Similar structures have been
demonstrated in the lower cryptogamic
plants (Fig. 57 c), but they appear to
be wanting in the cells of the higher
31
Cryptogams and the Phanerogams ( ).
The nucleus and cytoplasm are
the two most essential constituents
of the cell, and its vital functions de-

pend on the interaction between them.

In the lowest plants (Cyanophyceaeand KIU. 57. A nucleus <>f u c.-u (l f tu.- ymn-
""""-; B " ",
1>la t "' x JU 1 s" "
:

Bacteria)' such a division of labour in

.
",
'.'""
weed. '//, llio nttToandiBg cytoplasm ;
,
the protoplasm IS not certainly proved, ki the nucleus kw, nuclear membrane
; ;

the existence of a nucleus being still ". nucieoius; ceutrosome; a,, dn-o-
.-,

e j- /Q.}\ /~ir matoi>hores.

(x 1000.)
a matter of dispute ("). Cnroma-
tophores" are wanting in the Bacteria and Fungi, as in all animal
cells.
While animal usually remain continuously filled with
cells

protoplasm, vegetable soon form large SAP CAVITIES.

cells It is only
the embryonic cells of plants that are entirely filled with protoplasm,
as the cells, for example, of an embryo or of a growing point ; they
afterwards become larger and contain proportionally less protoplasm.
This can be seen in any longitudinal section through a stem apex.
At a short distance from the growing point the enlarged cells have
already begun to show cavities or VACUOLES (v in A, Fig. 58) in
their cytoplasm. These are filled with xa watery fluid, the CELL SAP.
The cells continue to increase in sie, and usually soon attain a
condition in which their whole central portion is filled by a single,
 BOTANY

large sap cavity (v in Fig. 58). The cytoplasm then forms only
,

a thin layer lining the cell wall, while the nucleus takes a parietal
position in the peripheral cytoplasmic layer.
At other times, however, the sap cavity of
a fully-developed cell may be traversed by
bands and threads of cytoplasm ; and in that
case the nucleus is suspended in the centre of
the But whatever position the nucleus
cell.

may occupy, it is always embedded in cyto-

plasm and there is always a continuous peri-
;

pheral layer of cytoplasm lining the cell wall.

This cytoplasmic peripheral layer is in con-
tact with the cell wall at all points, and, so
long as the remains living, it continues
cell
in that condition. In old cells, however, it
frequently becomes so thin as to escape
direct observation, and is not perceptible
until some dehydrating reagent, which causes
it to recede from the wall, has been employed.

Such a thin cytoplasmic peripheral layer has

been described by HUGO v. MOHL under the
name of PRIMORDIAL UTRICLE.
Dead cells lose their living
protoplasmic
contents, and, strictly speaking, should no
longer be termed cells, although the name
was first applied to them when in that, con-
dition. In reality they represent only cell
cavities. With their death, however, cells
do not lose their importance to a plant.
Without such cell cavities a highly-organised

plant could not exist, as they perform for

it the office of water-carriers, and afford
mechanical support and rigidity. The heart
wood of a tree consists exclusively of the
walls of dead cells.
The Protoplasm. In order to facilitate

Fio. 08. Two cells

.

taken at
an insight into the
real character of proto-
J
,
different distances from the plasm, attention will first be directed to the
growing point of a phanero- SLIME FUNGI
(Myxomycetes),a grOUpof Orgail-
gamic shoot. k. Nucleus ;. i_-i_ A.-UJUJ
it.
1SmS whlch stand On the b rder between the
!*, cytoplasm; v vacuoles,
, .

represented in B by the sap animal and vegetable kingdoms. These

cavity. (Somewhat diagram. are characterised at one stage
niatic, x circa 500.)
Myxomvcetes
J J , ,
, . . .1 t , t
of their development by the formation of a
PLASMODIUM, a large naked mass of protoplasm.
The plasmodium is formed from the protoplasm of the spores.
These spores are unicellular bodies (Fig. 59 a, fc), filled with cytoplasm,
 MORPHOLOGY 55

in which liesa central nucleus, and are surrounded by resistent cell

walls. The spores germinate in water, their contents, breaking through
the spore walls, come out (c, d) and round themselves off. change A
of form soon takes place ; the protoplasmic mass elongates and assumes
somewhat the shape of a pear, with the forward end prolonged into a

FIG. W.Clwtuli-ioderma di/ornte.. a, Dry, shrivelled spore; l>. swollen spore; c and </, s[>oics
showing escaping contents ; , /, g, swarm-spore changing to a myxoamoeba
swarm-spores ; h, ;

i, younger, k, older myxoamoeba; I, myxoamoebae about to fuse; m, small plasmodiuiu H, :

iwrtion of fully-developed plasmodium. ('<-, x 540: n, x 90.)

fine whip-like process or flagellum

(e, f, g).
Thus the contents of the
spore have become transformed into a SWARM-SPORE, which now swims
away by means of whip-like movements of its flagellum. In addition
to the nucleus, which is visible in the anterior end of every swarm-spore,
a vesicle may be seen at the other end, which, after gradually increas-
ing in size, suddenly vanishes, only to swell again into view. This
56 BOTANY I-AKT i

vesicle is a CONTRACTILE VACUOLE. The presence of such a contractile

vacuole in an organism was formerly considered a certain indication
of its animal nature. Xow, however, contractile vacuoles have been
observed in the swarm-spores of many green Algae, of whose vegetable
nature there can be no doubt.
The swarm-spores of the Myxomycetes soon lose this characteristic
swarm-movement, draw in their flagella, and pass into the amoeba
stage of their development, in which, like animal amoebae, they assume
irregular, constantly changing shapes, and are capable of performing
only amoeboid creeping movements. In the case of ChondriodermQ
tlifforme, a Myxomycete of frequent occurrence in rotting parts of
plants (Fig. 59), a number of the amoebae eventually collect together (/)
and coalesce. In this way, as is also the case with most other Myxo-
mycetes, the amoebae ultimately give rise to a plasmodium (n).
Although each one of the amoebae is so small that it can only be
seen with the aid of a microscope, the plasmodium into which they
become united may attain a considerable size.
The cytoplasm, both of the single amoeba and of the plasmodium,
consists of a clear ground substance, through which granules are dis-
tributed. This substance is of the consistence of a tenacious fluid ;
its superficial region is denser and free from granules, while these are
numerous in the less dense central portion. The granules enable the
internal streaming movements of the cytoplasm to be recognised.
The currents are constantly changing their direction, moving either
towards or away from the margin. The formation and withdrawal
of processes of the margin stand in relation to the direction of the
currents. When naked masses of protoplasm such as these plasmodia
encounter foreign bodies, they can enclose them in vacuoles, and,
when of use as food, digest them.
Deprived of its component water the protoplasm becomes hard and
tenacious, and, without losing its vitality, ceases to perform any of
its vital functions until again awakened into activity by a fresh supply

of water. In case of a scarcity of water the plasmodia of the Myxo-

mycetes may form SCLEROTIA, that is, masses of resting protoplasm of
an almost wax-like consistency. Months and indeed sometimes years
afterwards, it is possible for such sclerotia, if water be supplied, to
again produce motile plasmodia. Similarly, in seeds kept for a long
time, the protoplasm consolidates into a hard mass, which may be
easily cut with a knife, while the nuclei will be found to have shrunk
and lost their original shape. Nevertheless the protoplasts, after
absorbing water, may return again to a condition of activity.
Protoplasm is not a simple substance chemically ; it consists rather
of numerous different components, which are subject to continual

change. Since albuminous substances are always present, protoplasm

always gives a proteid reaction when incinerated, fumes of ammonia
;

are given off.

 MORPHOLOGY 57

Active protoplasm generally gives an alkaline, and, under certain conditions, a

neutral reaction, but never an acid one. The protoplasm of the higher plants
3
coagulates at a temperature not much over 50 C., in the Sehizophyta, however,
usually not below 75 C. In the inactive dried condition, as in spores and seeds,
it can endure a still higher temperature without coagulating. The spores of
U
many Bacteria can withstand a temperature as high as 105 C. Treated with
alcohol or ether, with acids of definite concentrations, with bichromates of the
alkali metals, or with corrosive sublimate, protoplasm quickly coagulates, while at
the same time insoluble proteid compounds are formed. Coagulating reagents,
accordingly, play an important part in microscopic technique ; of especial value
are those,' which, while fixing and hardening the protoplasm, change its structure
in the least degree. As fixing and hardening reagents for vegetable tissues,
alcohol, 1 per cent chromic acid, 1 per cent acetic acid, 0'5 to 1 per cent osmic
acid, concentrated picric acid, or corresponding mixtures of these acids, solutions of
mercuric chloride and formaldehyde, are used. Iodine stains protoplasm brownish
yellow nitric acid, followed by caustic potash, yellowish brown (xanthoprotein
;

reaction) sulphuric acid, if sugar be present, rose red.

;
Acid nitrate of mercury
(MiLLON's reagent) gives to protoplasm a brick-red colour. These reactions occur
with all proteid substances though they are not absolutely distinctive of them.
Protoplasm is soluble in dilute caustic potash and also in eau de Javelle (potassium-
hypochlorite), and accordingly both of these reagents may be recommended for
clearing specimens, when the cell contents are not to be investigated. All of the
above-mentioned reagents kill protoplasm until they have done so, their char-
;

acteristic reactions are not manifested. A large number of albuminous bodies or

.

albuminates have been named which are said to enter into the composition of
living protoplasm. Most of these compounds are still ill-defined ; in nuclei the
nucleins are most important, but they are also found in the cytoplasm. They are
characterised by containing much phosphorus, and are not attacked by pepsin,
and only with difficulty by trypsin. Staining reagents have also become an
important help to microscopic investigations for determining the composition of
protoplasm. This is due to the fact that the different constituents of protoplasm
take up and retain the stain with different degrees of intensity and energy. As a
general rule, only coagulated protoplasm can absorb colouring matter, although
some few aniline stains can, to a limited extent, permeate living protoplasts. For
staining vegetable protoplasts, which have been previously fixed, the various
carmines, hcematoxylin, safranin, iodine green, acid fuchsin, eosin, methylene
blue, and aniline bhie, gentian-violet and orange, have been found particularly
convenient. The different components of the protoplasm absorb the stains with
different intensities, and, when reagents are employed to remove the colouring
matters, they exhibit differences in their power to retain them. The nucleus
generally becomes more intensely coloured than the rest of the protoplasm,
especially a part of its substance, which is therefore called OHIIOMATIN. In
addition to those substances, which are to be regarded as integral parts of active
protoplasm, it always includes derivative products of albuininatcs, particularly
amides, such as asparagin, glutamin also ferments, such as diastase, pepsin,
;

invertin at times alkaloids, and always carbohydrates and fats.

;
The ash left
after incineration also shows that protoplasm always contains mineral matter,
even if only in small quantities. All such substances which do not enter
directly into the composition of protoplasm, but are only included within it, are

designated by the term METAPLASM.

The Cytoplasm. The cytoplasm of vegetable cells, which possess
58 BOTANY PART i

a cell wall, is a more or less tenacious fluid. It partakes of the

physical properties of fluids, and on being artificially freed from the

cell wall, tends to assume the spherical form. Its cohesion appears
to be greater in meristematic cells than in those which are older,
while in certain cases a still firmer consistence may be attained as
in the cilia borne by swarm-spores.
Both in the case of the Myxomycete and of the vegetable cell
enclosed by a wall, the basis of the cytoplasm consists of a hyaline
substance termed the HYALOPLASM. When granules are distributed
through the cytoplasm it is
spoken of as GRANULAR PLASMA or polio-
plasm. An extremely thin boundary layer is found at the periphery
which is quite free from granules, and a similar layer bounds
every vacuole present in the cytoplasm. The wall of the vacuole
is characterised by a greater
tenacity of life than the rest of the
cytoplasm, remaining alive for some time after the latter has been
killed by the action of a 10 per cent solution of potassium nitrate.
Since the vacuole- wall regulates the pressure exerted by the cell
sap contained in the vacuole, HUGO DE VRIES has applied the name
TONOPLAST (M) to this layer.
The small granules distributed through the granular plasma
consist of various substances, and may be classed together as MICRO-
SOMES. Some of them are small cavities filled with dissolved
substances,and to these the name PHYSODES has been given.
Even though bounded by a cell wall the cytoplasm frequently
exhibits movements comparable to those of the naked amoebae and
plasmodia of Myxonwcetes. These movements mostly are found in
somewhat old cells, but N. GAIDUKOV (^ has shown by means of
the recently invented ultramicroscope that they are of widespread
occurrence in vegetable protoplasts. The study of the movements
in the Myxomycetes showed that various kinds of movements could
be distinguished ; the waving movement of the flagellum of the
swarm-spore, the change in external form of amoebae and plasmodia,
to which their power of creeping about is due, and finally a streaming
movement in the cytoplasm. The cytoplasm, enclosed by a cell-
wall, may either exhibit isolated streaming movements, the direction
of which may undergo reversals, or a single stream, the direction of
which is constant. These two forms of movement are distinguished
as CIRCULATION and ROTATION respectively. In rotation, which is
found in cells with the cytoplasm reduced to a layer lining the
wall, the single continuous current follows the cell wall. In circula-
tion, on the other hand, the layer of cytoplasm lining the wall takes
no part in the movement, which is found in the strands traversing
the vacuole. Circulation common in cells of land-plants, while
is

rotation is more usual in

water-plants. The stimulus caused by
wounding the tissues in making the preparation frequently increases
the activity of the movement ( 35 ).
 MORPHOLOGY 59

A particularly favourable object for the study of protoplasm in circulation is

afforded by the staminal hairs of Tradescantia virginica. In each cell (Fig. 60)
small, fine currents of protoplasm flow in different directions in the peripheral
cytoplasmic layer, as well as in the cytoplasmic threads, which penetrate the sap
cavity. These cytoplasmic threads gradually change their form and structure, and
tli us alter the
position of the cell nucleus.
When the protoplasm is in rotation, the cell nucleus and chromatophores are
usually carried along by the current, but the chromatophores may remain in the
boundary layer, and thus not undergo movement. This is the case with the
Stoneworts (Characeae), whose long internodal cells, especially in the genus
NiteMa, afford good examples of well-marked rotation.

Properly fixed cytoplasm has a finely reticulate or honeycomb-

like structure, small granules being embedded in the network.
At particular developmental stages this structure appears to be
traversed by special filaments, which can be demonstrated by suitable
36
staining (Fig. 57) ( ).

The portion of the cytoplasm which forms the network or honeycomb appears
to be specially concerned with the nutritive processes, while the fibrillar plasma
influences the process of development they are respectively
;

termed trophoplasm and kinoplasm ;

the latter has also been
called archeplasm. When traced to their origins the limiting

layer of the cytoplast is found to belong to the kinoplasm,

the walls of vacuoles to the trophoplasm. Albuminous sub-
stances, precipitated by the fixing agent, have not infre-
quently been mistaken for structural features of the cyto-
s7
plasm ( ).

The Cell Nucleus ( ys ). The resting nucleus

has a reticulate or honeycomb structure forming
an anastomosing network (Fig. 56), which, how-
ever, in living objects can only be distinguished
by the punctated appearance it gives to the
nucleus. Streaming movements do not take place
within the nucleus. An insight into the nuclear
structure is only to be attained with the help of
properly fixed and stained preparations. It is then

possible to determine that the greater part of this

nuclear network is composed of delicate and, for
t:. CO. Cell from the most part, unstained threads, in which lie deeply
st.-uuinal hair of Tr
stained granules. The substance of the threads
intia virginicu,
'the'nucieus has been distinguished as LININ (I), that of the
by prato- granules as CHROMATIN (c/t). One or more larger
,,ias.,.ic strands, (x
bodies, the NUCLEOLI (n), occur at the intersections
of some of the linin threads ; these, although deeply
stained, do not take the same tint as the chromatin granules.
The network of the nucleus lies within the nuclear cavity, which is
r. i
BOTANY

filledwith nuclear sap and surrounded by a membrane (w). The

nuclear membrane, strictly speaking, is a part of the surrounding
cytoplasm, and is the protoplasmic layer with which the cytoplasm
separates itself from the nuclear cavity. The nucleus in young cells
with abundant protoplasm is, as a rule, spherical. AVhen situated in
the lining layer of cytoplasm of older cells, it is
frequently of a
flattened form, while
in elongated cells it

exhibits a correspond-
ing elongation. Ex-
ceptionally in old
cells the nucleus is

forke,., lobed, or of
some other irregular
shape. These changes
in form of the nucleus
are due to slow move-
ments, which cannot,
as a rule, be directly
observed. In em-
bryonic tissues the
nuclei are relatively
large in proportion
to the size of the pro-
Hie. 0-2. Portions i.r two al-

toplasts. Glandular .iacfnt cells in a hypha trnrn

cells are also t in' stalk lit' ;w/;.
usually .Nuclei
, : i..
pits.
provided with large (X o40.)
nuclei.
While the cells of the Cormophytes are
almost always uninucleate, in the Thallo-
phytes, on the contrary, multinucleate cells
are by no means infrequent. In the Fungi,
and in the
Siphoneae among the Algae, they
are the The whole plant is then
rule.

composed either of but one single multi-

,,,,
, extensively
chromic acid and stained with branched (Fig. 295), or it may consist of a
Nuclei
large number of multinuclear cells, forming
.aniline. n , ; ch, ciiro-
matophores ; p. pvrenoids; m, ..
,-> , .
,

stan-h grains. together one organism.

< X S40.) Thus, on suitable
treatment, several nuclei may be detected in
the peripheral cytoplasm of each of the cells of the common fila-
mentous fresh -water Alga Cladophora ylomerata (Fig. 6, p. 12,
Fig. 61).
The nuclei of the long, multinucleate cells (Fig. 62 n) of fungal
filaments, or HYPHyE, and also of many Siphoneae, are characterised
by their diminutive size.
 MORPHOLOGY til

The Centrioles and Centrosomes. In a number of cases among

the lower Cryptogams (Thallophyta and Bryophyta) structures which
correspond to the centrioles and centrosomes of animal cells have
been demonstrated. Where centrioles are found, as in some marine
Algae (Fucus, Fig. 57), they are dumb-bell shaped. Larger masses of
active cytoplasm occur in a number of Fungi, and, though they do
not, like them, include centrioles, are comparable with the centro-
somes of animal cells. Similar structures occur in the mother-cells
of the spermatozoids in Bryophyta, Pteridophyta, and certain Gym-
39
nosperms ( are there termed blepharoplasts, and provide
). They
the material for the formation of the cilia of the spermatozoids.
With the exception of the blepharoplasts no structures comparable
.with centrioles or centrosomes have been shown to exist in the cells
of the higher plants.
The Chromatophores ( ). In the embryonic cells of the embryo
4(l

and of growing points, where the chromatophores (Fig. 56 ch) are

principally located around the nucleus, they first appear as small,
colourless, highly refractive bodies. They may retain the same*
appearance in older cells (Fig. Ill A, I), but in them they also
attain a further development, as CHLOROPLASTS, LEUCOPLASTS, or
CHROMOPLASTS. Since these bodies have the same origin they are
all included in the one term, CHROMA TOPHORKS.

Chloroplasts. In parts of plants which are exposed to the

light the chromatophores usually develop into
chlorophyll bodies or chloroplasts. These are
generally green granules of a somewhat flat-
tened ellipsoidal shape (Fig. 63), and are scattered,
in numbers, in the parietal cytoplasm of the
cells. All the chloroplasts in the Cormophytes,
and, for the most part also, in the green
Thallophytes, have this form. In the lower
Algae, however, the chlorophyll bodies may
assume a band-like (Fig. 264 e), stellate or
tabular shape.

In these cases the chloroplast often includes one or more

pyrenoids ;
these are spherical protoplasmic bodies con-
taining an albuminous crystalloid, and are surrounded
by small grains of starch (Fig. 264). The ground sub-
stance of the chlorophyll bodies is itself colourless, but
contains numerous coloured granules, which are termed
:. ".'!. Two cells from ;i

leaf of Funaria liygi'n-

OKAXA. These consist of an oleaginous substance, which metrica. cl, Chlompla.st.s;
holds in solution green and yellow pigments. These a. nucleus, (x 300.)
colouring substances may be extracted by means of
alcohol, leaving only the colourless plasmic substance of the chlorophyll body
remaining.
The easiest
way which a solution of chlorophyll can be prepared, is to extract
in
the chlorophyll by means of alcohol from green -leaves that have been
previously
62 BOTANY PART i

boiled in water. The green chlorophyll pigment is also soluble in ether, fatty
and ethereal oils, paraffin, petroleum, and carbon disulphide. The alcoholic
solutions are fluorescent and appear green in transmitted light, blood red in
reflected light. When an alcoholic solution of chlorophyll is shaken up with
benzole, the latter, as KKAUS first showed, on standing rises to the surface as

a green solution, leaving the alcohol yellow.

41
According to MAHCHLEWSKI and ScHUNCK( ), two green pigments are to be
The one, true chlorophyll, is always
detected in the alcoholic extract from leaves.
present the other, which has been termed allochlorophyll, cannot be detected in
;

some plants. The yellow pigments of the chloroplasts are collectively termed
xanthophyll. Only chrysophyll, which forms shining red crystals, has been
isolated the rest of the xanthophyll form amorphous masses.
;
True chlorophyll
is by three absorption b;inds in the less refrangible half of the
characterised
spectrum, and three in the more refrangible portion. As yet only one band,
which lies in the red portion of the spectrum, is known for aliochlorophyll.
The amount of chlorophyll in a green plant is very small. T.SCHIRCH ( 4 -) has
calculated that only O'2-l'O gr. of chlorophyll can be obtained from a square-
metre of green foliage leaves.

From the investigations of MARCHLEWSKI and NENCKI it appears

that a relationship exists between chlorophyll and haemoglobin (the
43
pigment contained in red blood corpuscles) ( ).
The green colour of the chlorophyll in some groups of Algae is
more or less masked by other pigments. In addition to the chloro-
phyll green, with its accompanying yellow pigments, many of the
blue-green Schizophyceae contain a blue colouring matter, phycocyanin,
while the red Algae possess a red pigment termed phycoerythrin.
These pigments are soluble in water, and are characterised by a
beautiful fluorescence. The phycocyanin may often be found as a
blue border surrounding a blue-green Fission-Alga which has been
dried in a press.
Whether these pigments are mixed with the chlorophyll or are
44
chemically combined with it is as yet undecided ( ). Recently HANS
45
MOLISCH ( ) has attempted to prove that the brown coloration of the
Diatoms, the brown Algae, and especially of a saprophytic Orchid
(Neottia nidus a?vis), is not due to a mixture of a brown pigment with
chlorophyll. He regards it as due on the other hand to a single
pigment, phaeophyll, which is nearly related to chlorophyll and
readily undergoes a chemical change into ordinary chlorophyll.
Before the leaves of trees fall in the autumn their cells lose nearly
all their cytoplasmic contents, and at the same time the chloroplasts

undergo disorganisation. There remains only a watery substance in

the cell cavity, in which a few oil globules and crystals, together with
a few yellow, strongly refractive bodies, can be seen. Sometimes in
presence of abundant sugar this liquid in the cell cavities becomes red,
and thus imparts to the foliage its autumnal brilliancy. In the leaves
of coniferous trees, which only indicate the approaching winter by

assuming a somewhat brownish tint, the cause is different. The

SECT. I MORPHOLOGY 63

chlorophyll-green of their chloroplasts changes to a brownish green

pigment, but in the following spring regains its
characteristic colour.
In such phanerogamic parasites or humus-
plants as are devoid of green colour, the chloro-
plasts either do not develop, or they are white,
or have only a brownish or reddish colour. No
chromatophores are found in the Fungi.
Leueoplasts. In the interior of plants,
where light cannot penetrate, leucoplasts are de-
veloped from the rudiments of the chromato-
phores instead of chloroplasts. They are of a
denser consistency than the chloroplasts, mostly
spherical in shape, but often somewhat elongated
in consequence of enclosed albuminous crystals.
If the leucoplasts become at any time exposed to
FIG. 64. Cell from the
the light, they may change into chloroplasts. upper surface of the
This frequently occurs, for example, in the super- calyx of Trapaeolum
majus, showing chromo-
ficial portions of potato tubers. ]. lasts, (x 040.)
Chromoplasts. The chromoplasts most of
flowers and fruits arise either directly from the rudiments of
colourless chromatophores, or from previously formed chloroplasts.
In shape the chromoplasts resemble the chloroplasts, except that
they are usually smaller ;
in consequence of the crystallisation
of their colouring pigment they sometimes
assume a triangular, tabular, needle, or fan-
shaped form (Figs. 64, 65). The colour of the
chromoplasts varies from yellow to red, accord-
ing to the predominance of yellow xanthophyll
or orange -red carotin. The name carotin has
been derived from the Carrot (Daucus Carota),
in the roots of which it is particularly abundant

(Fig. 65). The frequent crystalline form of the

chromoplasts is, in a great part, due to the
tendency of carotin to crystallisation, although
itmay be also occasioned by needle-like crystals
of albumin. Xanthophyll, however, is never
KIG. Go. Cliromoplasts ,, f present in the chromoplasts except in an amor-
the Carrot, some with phous condition.
included starch grains.
Carotin is practically identical with the
(X 540.)
chrysophyll found in the chloroplasts. Its

spectrum only differs from that of chrysophyll in having the absorp-

tion bands slightly displaced towards the violet end ( 46 ).
47
Origin and Structure of the Cell Wall ( ). The membrane which
encloses the vegetable protoplast is a product of the protoplasm.
Many low organisms belonging to the Algae liberate naked protoplasts
64 BOTANY PART I

from their cells ; these swarm-spores (Fig. 97 A) serve to

multiply
the plant vegetatively. They soon settle down, form a thin cell
membrane on the surface of the protoplast, and proceed to give rise
to a filament. In more highly organised plants the ovum, from which
the development starts, has no cell wall until it has been fertilised ;

from this stage on, all the cells composing the plant are surrounded by
cell walls. At the growing points of plants the cells are separated
from one another only by extremely thin membranes or cell walls.
As the cells increase in number by repeated division, new cell walls
are being continually introduced between the existing ones. The

l-'ic. lii. Part, of asclerenchj-

FIG. thickened cell inatous libre IKnii Vln<:
t>G.--Htrongly
from the pith of Cli'indtix rHnllm. iiiftjm: The striationsof the
Middle lamella intercellular outer layer* are in<>re appar-
,
. ; i,
ent than those of the inner
space; pit; ie, pitted} cell wall
t,

in surface view. (X 300.) layers. The thicknessof the

wall, as seen in optical sec-
tion, is also .shown. (X500.)

rapid growth in length which sets in a short distance from the

growing point, as a result of the increase in the size of the cells, must
be accompanied by a corresponding GROWTH IN SURFACE of the cell
walls. So long as this growth in surface continues, the cell walls
remain thin. After the cells have attained their ultimate size, the
GROWTH IN THICKNESS of the cell walls then begins. The growth in
surface of the cell wall may either involve the introduction of new
material, ormay take place without this. In the latter case the mem-
brane would become thinner if new lamellae were not simultaneously
applied to its surface. Growth of the wall by the introduction of
new particles between those previously existing is termed GROWTH BY
INTUSSUSCEPTION, while that which occurs by the laying down of new
lamellae on the surface of the older ones is called GROWTH BY APPOSI-
 MORPHOLOGY . 65

TION. The later growth in thickness of most cell walls takes place by
apposition, and thus the stratification, Avhich such thickened walls
exhibit, is brought about (Fig. 66). Thicker, dense layers alternate
with thinner less dense ones. The denser
layers can be recognised by their high re-
fractive power. In many cases lamella,
deposited by apposition, become further
thickened and otherwise modified by a
process of intussusception. Three distinct
layers can frequently be distinguished in
strongly thickened cell walls, such as those
of the wood, a primary, a secondary, and
a tertiary thickening layer these differ in;

their optical appearance and their chemical

composition. The secondary thickening

layer is usually the most strongly devel-
oped, and forms the chief part of the cell
wall. The tertiary or inner layer is usually
more highly refractive.

Fl 6s. -Surface view of cells from

Thicker cell walls or layers of the wall which -

the sensitive side of the tendril

appear homogeneous frequently exhibit a stratifica- of Cucurbita Pepo< showill , ta ,.
tion when treated with strong acids or alkalies. In tile pits, s. (x 540.)
many cases the thickening layers exhibit delicate
striations in surface view. The striations extend through the whole thickness of
the layers, usually running obliquely to the long axis of the cell, and often crossing
one another in the different thickening layers (Fig. 67). A similar appearance of
crossing of striations may result from the single thickening layers of two adjoining
cells being visible at once. In special cases, but only in the formation of repro-
ductive cells, an inner thickening layer, completely
detached from the others, is produced, as in the
formation of pollen grains and the spores of
Bryophytes and Pteridophytes, which, enclosed
only by this inner membrance, finally become
freed from the older thickening layer. This process
is often alluded to as REJUVENESCENCE in such ;

cases, it should be noted, there are, in reality, no

new cells formed.
Bio. 69. T,a,,sv,rs, section thro^h
similar ceils to those in Fi K 08; a
-phe thickening of the cell wall seldom
.

small crystal of calcium oxalate is

, , < i ,111 <

present
takes
in thetactiiepits.
(x4so.) place uniformly over the whole surf ace;
but some portions are thickened, while, at
other points, the original cell wall remains thin. In this way pores
are formed which penetrate the thickening layers. These pores
or PITS may be either circular (Figs. 66 w, 71 m), elliptical, or
elongated. The pits in adjoining cells coincide, and would form one
continuous canal, were not that the unthickened primary cell wall
it

persists as a PIT MEMBRANE between the two pits. The openings

of narrow elliptical pits into adjoining cells usually appear to cross
F
66 BOTANY

one another obliquely. As a result of the continued thickening of

6'"'

FIG. 70. From the wood of the Pine, 7'i/itts sylvextris.

A, Hordered pit in surface view ;B, bordered pit in

tiingential section; t, torus; '', transverse section of
Ki<;. 71. Cells from the endo-
a tracheid ; m, middle lamella, with gusset, M* ;

sperm of Hr/iUhoiial"-'
t, inner peripheral layer, (x Mil.)
In Hutu in. >[/. I'its in surface
view ; p, closing ineiiilirane ;

it, liueleus. (X -40.)

the cell wall, the canals of several pits often unite, and so BRANCH KD
PITS are formed. Such branched pits have usually very narrow

A
Fio. 72. Part of two KK.. ~X. A, Part of ;in anniil;ir trnrheid ;

sieve -
tuU's of the ]:. ]>art of a s]>ir:d trai-heiil ;
C, l"ir^i-

Pine, I'iiniK xi/l'-

tudinal section through |<iirt of a

showint; sieve -
pits.
retic'iil;ite vessel, showing: the remains
(x 540.) of :i
partition wall, x. ( 1MH.)

canals, and occur for the most part only in extremely thick and hard
cell walls, as, for instance, in those of the so-called sclerotic cells or
 67

sclereides. Simple pits may, on the other hand, expand on

approaching the primary cell wall.
Pits widened towards the membrane are found in the external
cell walls of many tendrils 4S These pits, which are filled with cyto-
( ).

plasm, probably receive

the stimulus, and may
be termed tactile pits
(Figs. 68, 69). The
structures known as
BORDERED PITS (Fig. 70)
are a special type of
simple pits widened to-
wards the pit membrane.
The pit may be present
on one or both sides of
the wall; the former is
the case when the water-
conducting element
abuts on a cell with Km. 74. Purl of transverse section of a stem of Inipatien.*
pai-viflora. e. Epidermis ; c, colleiicliynia ; )>, tliin-walk'<l
protoplasmic contents, parenchymatuus ct-Ils intercellular sjwce. (x 300.)
; i,

the latter when the

pitted wall separates two water-conducting elements. In bordered
pits the closing membrane is thickened at the centre to form a TORUS
(Fig. 70, B). By the curving of the closing membrane to one side or
the other the torus may so act as to close the pit canal (Fig, 70 P>, f).
The bordered pits apparently act as valves. Seen from the surface a
bordered pit appears as two con-
centric rings (Fig. 70 A). The
smaller, inner ring represents the
narrow opening of the pit into the
cell cavity ; the larger, outer ring
indicates the widest portion of the
PIT CHAMBER when it abuts on the
primary cell wall.
The pit membrane of specially
Pto. 75.- Epidermal cell from the margin of a ., .
,, , ., , .

r-uiicai leaf of a,,,,,,,,,,,,!,, ,..,*;,//./,. The wide P8

between cells with thin
lens-shaped thickening of the outer wall is walls often shows thicker bands
in this plant silicitied. (After HABERLANDT,
X
wh j ch g j ye i(
.

ft l att j ce _ wor k appear-

515.)
A
similar type of thickening
ance.
is found in the membranes of the sieve-pits which, on account of
their being perforated like a sieve, are termed sieve-plates (Fig. 72).
In cases where the greater part of the cell wall remains un-
thickened, its character is determined by its thickened rather than

by unthickened
its portions ; it is in this sense that the terms
annular, spiral, and reticulate are used (Fig. 73). Just as in the case
of cells with bordered pits, annular, spiral, and reticulate cell walls
68 BOTANY I'AUT I

are only acquired by cells that soon lose their contents, and act as
water-carriers. Such wall thickenings serve as mechanical supports,
to give rigidity to the cells, and to
enable the cell walls to withstand
the pressure of the surrounding
living cells.

The thickened bands by which

thickenings of the cell wall are effected
an: attached by narrowed bases to the

primary membrane (
49
). When the mem-
Inane separates two water - conducting
elements its central portion is thickened
like the torus of the bordered pit. The
annular and spiral types of thicki-niiii:
characterise the water -conducting ele-
ments of growing parts of the plant,
>ince they allow of extension of the wall

iluring growth. The thickening band

can often be removed as a continuous
spiral from the lamella to which it i*

attached.

COLLENCHYMATOUS
cells are
Fu, 7G.-l>art of transverse
Fieus dastica. c, Cystoiitu e,
section of a leaf of
e, three-
;
lg
the Walls of which are
cells,
thickened principally at the corners
,

layered epidermis: ;. palisade ^renchyma \

?
/pj g 74 c suc ^ ce ]J g occur com .
:

*, spongy parenchyma, (x '-'40.) , /, , , , .

, . .

monly in the more highly organised

plants and form a special kind of tissue (p. 116). Cells on the
surface of plants have usually only their outer walls thickened
,

(Fig. 74 e).

Unequal thickening of the cell- wall

is most noticeable when it is limited
This is the case
to definite small areas.
with those lens-shaped thickenings of
the outer walls of the epidermal cells
of the upper surface of foliage leaves,
which are of frequent occurrence and,
to G. HABERLANPT M ),
according (

act as convex lenses to concentrate

the light (Fig. 75). By the thicken-
ing of cell walls at special points,
protuberances projecting into the cell
1

Kn;. 77. -.1, I'nlleii-s.'i-ain nt Cucurbita I '/-" iii .sin--

cavity are formed ;

in this way the face view, and partly in uptiral sertiiin. rendered
formations known as CYSTOLITHS transparent I'.v treating with oil of lemons

arise. Certain large cells in the leaves (x 240); R, part of transver-r src-tinn of pollen

.urain of Cifurliittt re rnif'Sa ( x ">4<i).

of the India-rubber plant (Ficus elas-
tica)contain peculiar clustered bodies, formed by the thickening of the cell wall
at a single pointj (Fig. 76). In their formation a stem-like body or stalk first
protrudes from the cell wall ; by the addition of freshly deposited layers this
SECT, i MORPHOLOGY 69

becomes club-shaped, and, by continued irregular deposits, it finally attains its

clustered form.

So only centripetal wall thickenings have been described.

far
Cells, the walls of which are centrifugally thickened, can naturally
only occur where the cell walls have free surfaces. The outer walls
of hairs generally show small inequalities and projections. The
surface walls of spores and pollen grains (Fig. 77) show a great
variety of such centrifugally developed protuberances, in the form of
spines, ridges, reticulations, and bands of characteristic structure.
51
Cell-Wall Substance ( ).
The most important constituent of
cell Avails is CELLULOSE.
It is present in the cell walls of all plants,

except in those of the majority of Fungi.

Cellulose is a carbohydrate of which the chemical composition is expressed by
the general formula (C 6 10 5 )n. H
It is insoluble in dilute acids or alkalies. By
the action of concentrated sulphuric acid it is converted into dextrose. After
treatment with sulphuric or phosphoric acid, iodine will colour it blue it shows a ;

similar reaction when exposed to the simultaneous action of a concentrated solution

of certain salts, such as zinc chloride or aluminium chloride, and of iodine.
Accordingly, chlor-zinc-iodide, on account of the blue colour imparted by it, is
one of the most convenient tests for cellulose. GILSON ( M ) obtained cellulose in a
crystalline condition in the form of sphaerites or dendrites.

The walls never consist entirely of pure cellulose, but contain

cell
a considerable amount of other substances, which are not stained blue
by chloroiodide of zinc. In unlignified cell walls PECTIN SUBSTANCES
are particularly prominent. They are easily distinguished by the
readiness with which they dissolve in alkalies, after being previously
acted upon by a dilute acid.
for example, congo red
Susceptibility to certain stains is a characteristic of

cellulose while other stains, such as safranin and methylene blue, colour pectin
;

M
substances more deeply. According to MANGIX ( ), the partition wall formed in
the higher plants during cell division consists almost wholly of pectin substances ;

the next developed laminae, the secondary cell- wall layer, of a mixture of cellulose
and pectin substances the last formed, or tertiary layer, chiefly of cellulose. If
;

the secondary layer of the cell wall remain unlignified, the amount of the pectin
substances contained in it increases with age and helps to strengthen the MIDDLE
i,
\MKLLA, or primary cell- wall layer.

Amongthe substances entering into the composition of cell walls, in addition to

cellulose and the pectin substances, mention must be made of CALLOSE. It is

characterised by its insolubility in cuprammonia and solubility in soda solution,

and in a cold 1 per cent solution of caustic potash. It is coloured a red brown by
chloroiodide of zinc, with aniline blue it takes an intense blue, and with corallin
(rosolic acid) a brilliant red. Its presence in the higher plants is limited to a few

special cases it envelops the sieve-pits and is always present in calcified cell-wall
;

layers, as, example, in cystoliths (Fig. 76). Chitin, a proteid substance,

for
M
according to GILSON ( ), takes the same place in the cell walls of the Fungi as
cellulose in the cell walls of the higher plants.

Where cell walls become LIGNIFIED or SUBERISED, it is

particularly
70 BOTANY I-AKT i

the secondary layer that receives the wood or cork substance, while
the tertiary or internal layer retains its cellulose character.

Lignificatiou depends on the introduction into the membrane of certain

substances, among which, according to CZAPEK an aromatic aldehyde which
(**),

he names hadromal is never wanting. According to F. C. vox FAIJEU ("*) hadromal

is not present in all lignified membranes, and on the other hand occurs in many

that are not lignified. Associated with hadromal, according to CZAPEK, is coniferin,
whieh can be obtained from the youngest xylem. C/AP-EK denies 1 1n-, existence of
vanillin in lignified membranes. The proportion in whicli hadromal is found in
wood does not exceed 1-2 per cent of its dry weight. To its presence the so-callcil
liguin reactions are due, a violet colouration with phloroglucin and hydrochloric
acid, and a yellow colouration with acid anilin sulphate. The potassium per-
manganate reaction (a red colouration on treatment with a 1 per cent solution of
that substance followed by ammonia) is, according to F. C. FABER, a general feature
of lignified membranes. With chlor-zinc-iodide lignified membranes stain yellow,
not blue.
Corky cell walls contain suberin and take a yellowish-brown colour with chloro-
iodide of zinc with caustic potash, a yellow. VAN AVissF.r.iNiac ( s~) has disputed
;

the presence of cellulose in suberised cell walls, and regards the cork substance or
.sruKUix as a fatty body, which is composed of glycerine esters and other com-
pound esters, as well as of one or more other substances which are infusible, insoluble
in chloroform, and decomposed by a solution of caustic potash.

CUTINISATION, which is similar to but not identical with suberisa-

tion, is usually due to the subsequent deposition of cutin in cellulose
cell walls.

VAN has shown that phellonic acid, which is always present in

"\VI.SSKI.INGH

suberiu, constantly absent in cutin.

is The behaviour of cutin, as of suberin, varies
according to the source from whicli it is derived. Cutin withstands better the
action of- caustic potash. In other respects, the reactions given by cutinised cell
w alls with chloroiodide of
r
zinc or solutions of caustic potash are almost identical
with those of suberised cell walls.

Young cell walls are less elastic, but relatively more extensible
than older ones. The power of resisting a stress
is increased by

lignification. The presence and

of cutinised
corky membranes at the
surface of the plant diminishes the loss of water from it (M
The layers of the cell walls of some cells, particularly the super-
ficial cells of certain fruits, as of Sage, and of numerous seeds, such as
Flax and Quince seeds, become mucilaginous, ;tnd swell in water to
59
MUCILAGE, which, according to),
KLEBS
serves the purpose of
G. (

attaching the seeds to the soil. Firm

cell walls can also be trans
formed into GUM, as is so often apparent in Cherry and Acacia trees,
portions of the wood of which often succumb to GU.MMUSIS.

The several varieties of gvms and mucilage react differently, according as they
are derived from cellulose, callose, pectin substances, or from allied sulistam -e*.
According to MANC.IN they may be microchemically distinguished by their ivai-tit>n
with ruthenium red, which stains only such as are derived from pectin -nlt.un .-
SKCT. i MORPHOLOGY 71

or related substances, such as the mucilage of the seeds of the Cruciferae and Quince
(Cydonia), that from the mucilage cells of the Malveae, the gums of the Cherry
and Acacia, the gum tragacanth from Astragalus gummifer. The mucilage of
Orchid tubers, on the other hand, is related to cellulose, and remains uncoloured
with the same reagent.
The cell walls of the seeds of many Palms, as also those of Ornithogalum
(Fig. 71), have strongly developed thickening layers, which are full of pits. These
thickening layers are lustrous white, and, as in the case of the seeds of the Palm,
I'/iiff: Icphas macrocarpa, may attain such a degree of hardness as to be technically

valuable as vegetable ivory. Such thickening layers may contain other carbo-
hydrates in addition to cellulose, and by the action of ferments are dissolved
during germination. They are accordingly to be considered as a reserve substance
of the seeds.

Cell walls often become coloured by tannin or derivative sub-

stances ; way, for instance, the dark colour is produced which
in this
is often seen in the coats of seeds and in old wood. The colours of
the woods of economic value are due to such discoloured cell walls.
Inorganic substances are often deposited in large quantities in old cell
walls. Among such substances calcium oxalate is often met with,
commonly in crystal form ; also, although not so frequently, calcium
carbonate. In the cystoliths of Ficus elastica (Fig. 76) so much
calcium carbonate is deposited that it effervesces with hydrochloric
acid. In many plants, as, for instance, most of the Characeae, the
quantity of calcium carbonate in the cell walls is so great as to
render them stiff and brittle. Silica is also present in the superficial
cell walls of the Gramineae, Equisetaceae, and many other plants, and

gives them a very considerable firmness. The lens-shaped thickenings

in the outer walls of the epidermal cells of Campanula persicifolia are
also silicified.

By withdrawing water from the cells a contraction of the protoplast and its
consequent separation from the cell wall is brought about (see Plasmolysis). Such
protoplasts are able under certain conditions to surround themselves with a new
cell -membrane. The removal of the cuticle or of the waxy covering from the
surface of certain plants (Agave, Aloe, Ricinus, Sedum) is followed by its re
60
generation ( ).

Form of the Cell. As cytoplasm is a viscous fluid, and would tend,

ifunimpeded, to take a spherical shape, it may be assumed that the
natural and primary form for cells is spherical. Such a shape,
however, could only be realised by cells which, in their living
condition, were completely free and unconfined, or in such as were
able to expand freely in all directions. Newly developed cells,
in a continuous tissue, are, at first, nearly always polygonal.
Through subsequent growth their shape may change. The cubical
cellsof the growing point either elongate to a prism or remain,
owing to repeated division, short and tabular. If the growth is

limited to certain regularly arranged points of the surface, they

72 BOTANY

become stellate ;
if these points are less
uniformly arranged their
outline is unsymmetrical.
correspondingly In consequence of
energetic growth in length, fibre-like, pointed cells are developed.
If the walls of such cells become much thickened, they are called
SCLERENCHYMA fibres (Fig. 78 A}. These show diagonal markings,
due to their elongated pits, which are generally but few in number.
When fully developed, the living contents
A R L f suc h ce ^ s are sma U i amount and
frequently they contain only air. In the
last case, they merely act as mechanical
elements (stereides) and contribute to the
rigidity of the plant as a whole. Cells
somewhat similar,but shorter and consider-
ably wider, not pointed at the ends, and
provided with bordered pits, are called
TRACHEIDES (Fig. 78 B}. The tracheides,
in their fully developed condition, never
have any living contents, but serve as
water-carriers for the plant. So long as
they remain active, they contain only water
and isolated air -bubbles; their active
functions afterwards cease, and they become
filled with air. Tracheides which are speci-
ally elongated, and at the same time have
only a narrow lumen, and, like the scleren-
chymatous fibres, serve merely mechanical
purposes, are known as FIBRE TRACHEIDES.
Very long tracheides with a wide lumen
and thin walls, functioning, like typical
tracheids, as water-carriers, are distin-
guished as vasiform or VASCULAR TRACHE-
IDES (Fig. 78 C). They are characterised by
Fio. 78. .-I, A
sclerenchyinatous ,1 i i i' i i
the annular, spiral, or reticulate markings
tibre: js.atracheid: c, part of
a spiral tracheid ; D, part of a of their thickening layers, and may also be
latex tube. x loo
pr0 vided with bordered pits. The walls of
(A, K, c, ; D,

tracheides are always lignified, while those

of the sclerenchyma fibres may or may not have undergone this change.
Of all the cells in the more highly organised plants, the LATEX
CELLS or milk cells, also spoken of as latex tubes, attain the greatest
length. In the Euphorbiaceae, Moraceae, Apocynaceae, and Ascle-
piadaceae they arise from cells which are already differentiated in the
embryo. Growing as the embryo grows, they branch with it and
penetrate all its members, and may thus ultimately become many
metres long. The latex cells themselves have, for the most part,
unthickened, smooth, elastic Avails which give a cellulose reaction.
They are provided ( 01 ) with a peripheral layer of living cytoplasm
 MORPHOLOGY 73

and numerous nuclei. Their sap is a milky, usually white fluid,

which contains gum-resins, i.e. mixtures of gums and resins,
caoutchouc, fat and wax in emulsion. In addition, they sometimes
hold in solution enzymes, leptomin, tannins, often poisonous
alkaloids, and salts, especially calcium malate, also in the case of
Ficus Carica and Carica Papaya peptonising ferments. Proteid
granules often occur in the latex, and in the latex cells of the
Euphorbiaceae there are also present peculiar dumb-bell-shaped starch
grains. On exposure to the air the milky sap quickly coagulates.
In the adjoining figure (Fig. 78 D) is shown a portion of an isolated
latex cell dissected out of the stem of an Asclepiadaceous plant,
Ceropegia stapelioides.
Special cells, which differ in form, contents, or in their peculiar
wall thickenings from their neighbouring cells, are distinguished as
IDIOBLASTS. If strongly thickened and lignified, they are called
sclerotic cells (stone cells) or sclereides. They often contain ferments;
in the Cruciferae and some other orders myrosin is thus present,
Avhile Prunus laurocerasus contains emulsin. For the most part they
contain excreted substances such as tannins and calcium oxalate.
In Fig. 85 an idioblast, containing a bundle of raphides, is represented.
Idioblasts, resembling tracheides and functioning as water reservoirs,
are found between the chlorophyll-containing cells in the leaves of
some of the Orchidaceae and Cactaceae.
Size of Cells. The corresponding cells of equivalent members
of thesame plant are usually of nearly the same size, even when the
members show a variation in size.
Differentiation of the Protoplasts. In organisms composed
of one or of few cells the separate parts of the same protoplasts
may under some circumstances exhibit marked differences. On the
other hand, in multicellular organisms whole protoplasts are specialised
for functions with the performance of which their definite structure
is connected.
Inclusions Protoplasm STARCH. The chloroplasts in
of the
plants exposed the light almost always contain starch grains.
to
These grains of starch found in the chloroplasts are the first visible
products of the assimilation of inorganic matter. They are formed
in large numbers, but as they are continually dissolving, always
remain small. Large starch grains are found only in the reservoirs
of reserve material, where starch is formed from the deposited

products of previous assimilation. Such starch is termed RESERVE

STARCH, in contrast to the ASSIMILATION STARCH formed in the
chloroplasts. All starch used for economic purposes is reserve
starch. The starch grains stored as reserve material in potatoes are
comparatively large, attaining an average size of 0'09 mm. As
shown in the following figure (Fig. 79), they are plainly stratified.
The stratification is due to the varying densities of the successive
74 I'.OTAXY

layers; thicker denser layers which appear clear by transmitted

light alternate with thinner less dense layers which appear dark.
They are excentric in structure, since the organic centre, about which
the different layers are laid down, does not correspond with the centre
of the grain. The starch grains of the leguminous plants and cereals,

FIG. 70. Starch grains from a potato. A, simple ; KKI. So. Starch grains
.B, half-compound ; C and D, compound starch from the cotyledons <>f

grains c, organic centre of the starch grains, or

; Ph<i.--< -.
( x
nucleus of theii formation, (x MO.) 540.);

on the other hand, are concentric, and the nucleus of their formation
is in the centre of the grain. The
starch grains of the Kidney Bean,
Phaseohis vulfjaris (Fig. 80), have the shape of a flattened sphere or
ellipsoid ; they show a distinct stratification, and are crossed by
fissures radiating from the centre. The
disc-shaped starch grains of wheat are of un-
equal size, and only indistinctly stratified.
A comparison of the accompanying figures
(Figs. 79-81), all equally magnified, will give
an idea of the varying size of the starch grains
of different Pknts. The size of starch V*
pound grain; B, isolated varies, in fact, from 0'002 mm. to 0*170 mm.
grains 0'170 mm. large, such as those
comjionftnt grains of a com- Starch
pound grain, (x ">4o.)
from the rhizome of Canna, may be seen even
with the naked eye, as minute bright bodies. In addition to the simple
starch grains so far described, half-compound and compound starch
grains are often found. Grains of the former kind are made up of two
or more individual grains, surrounded by a zone of peripheral layers
enveloping them in common. The compound
grains consist merely
of an aggregate of individual grains unprovided with any common
enveloping layers. Both half-compound (Fig. 79 ) and compound
 MORPHOLOGY 75

starch grains (Fig. 79 C, D) occur in potatoes, together with simple

grains. In oats (Fig. 81) and rice all the starch grains are com-
62
pound. According to NAGELI ( ), the compound starch grains of
from 4 to 100 single grains; those of
rice consist of the oat of about
300, and those of Spinacia glabra sometimes of over 30,000. Starch
thus formed from previously assimilated organic substances also
requires chromatophores for its production. The grains are formed by
means of leucoplasts, which are, in consequence, often termed STARCH-
BUILDERS. If the starch grain is
uniformly surrounded by the
leucoplast during its formation it
grows uniformly on all sidesand
is
symmetrical about its centre. If
the formation of a starch grain should
begin near the periphery of a leuco-
plast, the grain would grow more
rnpicHy on the side on which the main
mass of the leucoplast is present and
the starch grain thus becomes excentric
(Fig. 82). Should, however, several
starch grains commence to form at the
same time in one leucoplast, they would
become crowded together and form
a compound starch grain, which, if ad-
ditional starchy layers are laid down,
gives rise to a half-compound grain.

It lias been asserted that starch grains are

crystalline bodies, so-called sphserites ( ), and
K! Fw. 82. LeaeoptaiU from an aerial tubwr
of plu 'J us ;/<"<''>' <'"<
are composed of fine, radially arranged, Viewed from the side ; /.', viewed from
needle -
shaped which A. MEYER
crystals, above leucoiilast becoming green
; /.',

terms Their stratification, accord-

trichites. and changing to a chloroplast. (x 540.)
ing to this view, is due to variations in the
form and number of the crystal needles in the successive layers, On the
other hand, H. FISCHER ( e4 ) has explained the stratification as due to zonal splits
rich in water, which originate by contraction taking place in the substance
of the grain at some distance from its growing surface. HENRY KRAEMER ( ) 6o

holds that a crystalloid and a colloid substance are present, but are united in
different proportions in the successive lamellae of the starch grain. In a few in-
dividual cases, ARTHUR MEYER has succeeded in showing that the stratification of
the starch grains corresponds to the alternation of the periods of day and night, i.e.
to the interference which is thus caused in the nutritive processes. The growth of
starch grains is also, according to him, affected by the solvent action of surrounding
substances, whereby the peripheral layers may be partially removed, and then no
longer completely envelop the entire grain. Starch grains are composed of a
carbohydrate, the formula of which is (C 6 H 10 3 )n. Most starch-grains consist of
amylose and are coloured blue with iodine in addition to this they contain, according
;

to L. MAQUEXXK and EUCJ. Roux C 66 ), a mucilaginous substance which they term

amylo-pectin. The presence of this substance causes the pasty union of the starch-
grains when treated with boiling water or alkalies. The grains swell in water of
3
60-70 C., are but little soluble in water at 100 C., but are rapidly dissolved by
 BOTANY

3
super-heated water at 140 -150 C. Starch swells very readily at ordinary
temperatures in solutions of potassium, or sodium hydrate. Heated without
addition of water, i.e. roasted, starch becomes transformed into dextrin, and is
then soluble in water and correspondingly more digestible. The glutinous starch
of varieties of Rice and Millet (Oryza saliva, var. ylntinoxx, Xr</hii>/i nil'jurc,
ylutinosum) is in the completely unswollen condition coloured brown with iodine ;

when swollen by a dilute solution of iodine, it takes a red, wine-red, or purple

m
colour. According to 0. BUTSCHLI ( ) it consists of amyloerythrin, a carbo-
hydrate resembling starch. In polarised light, starch grains, like inorganic
sphferites, show a dark cross. This appearance is usually referred to the doubly
refractive nature of the elements of which the starch grain is constructed.

The amount of starch contained in reservoirs of reserve material

is often considerable ; in the case of potatoes 25 per cent of their
whole weight is reserve starch, and in wheat the proportion of starch
is as high as 70 per cent. The starch flour of economic use is derived
by washing out the starch from such reservoirs of reserve starch. In
the preparation of ordinary flour, on the contrary, the tissues con-
taining the starch are retained in the process of milling.
Aleurone. Aleurone or proteid grains are produced in the seeds
of numerous plants, especially in those containing oil. They are
formed from vacuoles, the contents of which are rich in albumen,
and harden into round grains or, sometimes, into irregularly shaped
bodies. The albuminous substances of which they consist are, accord-
8
ing to A. TSCHIKGH and H. KRITZLER ( ), mainly globulins. A
portion of the albumen often crystallises, so that frequently one and
occasionally several crystals are formed within the aleurone grain.
In aleurone grains containing albumen crystals there may often be
found globular bodies, termed GLOBOIDS, which, according to PFEFFER
9
( ),
consist of a double phosphate of magnesium and calcium in
combination with some organic
A substances. Crystals of calcium
Jt
oxalate are also found enclosed
in aleurone grains.

ff The seeds of Ricinus (Fig. 83)

furnish good examples of aleurone
grains with enclosed albumen crystals
and globoids. The aleurone grains
themselves lie embedded in a cyto-
plasm that is rich in oil. In the
FIG. 83. A, Cell from the endosperm of Ricinus cereals the aleurone grains which lie
wnmttni*, in water; 1'. isolated aleurone grains
in olive oil albumen crystals a, glolioid.
Q^ .
JQ t j )e Quter
_. ,.
^ j
11
of the
i <
seeds (Fig. 84 at) are small, and free
; I:, ;

from all inclusions they

;
contain
neither crystals nor globoids. As the outer cells of wheat grains contain only
is the
aleurone, and the inner almost exclusively starch, it follows that flour
richer or poorer in albumen, the more or less completely this outer layer has been
removed before the wheat is ground. From the inner layers finer and whiter
 MORPHOLOGY 77

flour can be made while more nourishing Hour is obtained from the outer
:

layers. Reactions for aleurone are the same as those already mentioned for
the albuminous substance of protoplasm. Treatment of a cross-section of a grain
of wheat (Fig. 84) with a solution of iodine would give the aleurone layer a yellow-
brown colour, while the starch layers would be coloured blue.
ALBUMEN CUYSTALS. Crystals of albumen are of relatively frequent occurrence
in vegetable tissues and are often found in aleurone grains (Fig. 83). Their
appearance in the seed of Ricinus has been described above, and especially large
crystals are found in the endosperm of the Brazil nut (BertlmHrtiti. ov/Vs^ i

if

Ki<;. S4. Part of u section -rain ol' wheat, Tfitii-mn

M!';I

i-
nil/are, p, Pericarp ; t,seed coat, internal to which
is the endosperm; al, aleurone grains; mn, starch
grains ; //, cell nucleus, (x 240.)
KlO. 85. Cell I'r the cortex
of Drvaienti i-ubrn, tilled

with mucilaginous matter

and containing a bundle
of raphides. /. (x HiO.)

ing to the Lecythidaceae. They have previously been mentioned as occurring in

the chromatophores. In the illustration of the leucoplasts of Phajus grandifolius
(Fig. 82), the rod-shaped crystals are represented as light stripes (in and E). B
Albumen crystals may also occur directly in the cytoplasm ; as, for instance, in
the cells poor in starch, in the peripheral layers of potatoes. Albumen crystals
are sometimes found even in the cell nucleus. This is particularly the case in the
Toothwoi t (Lathraea squamaria], and in many Scrophulariaceae and Oleaceae ( 70 ).
Albumen crystals usually belong either to the regular or to the hexagonal crystal
system. They differ from other crystals in that, like dead albuminous substances,
they may be stained, and also in that they are capable of swelling by imbibition.
Subjected to the action of water or a dilute solution of caustic potash, they at first
increase in size without losing their crystalline outline.
?8 BOTANY I'AKT i

Crystals of Calcium Oxalate. Few plants are devoid of such

They are formed in the cytoplasm, within A acuoles Avhk-h
r
crystals.
afterwards enlarge and sometimes almost fill the whole cell. In such
cases the other components of the cell become greatly reduced; the
cell walls at the same time are often converted into cork, and the
whole cell becomes merely a repository for the crystal. The crystals
may be developed singly in a cell, in which case they are of consiiU-i
able size and can be seen to belong either to the tetragonal or
monosymmetrical crystal system ; or the crystals are so small and
numerous that their form cannot be clearly made out and they appear
as a crystalline sand filling the cell. Frequently they form CRYSTAL
AGGREGATES, Clusters of crystals radiating in all directions from a
common centre. In the Liliaceae, Orchidaceae, and other Mono-
cotyledons, compact bundles of needle-shaped crystals of calcium
oxalate, the so-called RAPHIDES, are especially frequent (Fig. 85).
Such crystal bundles are always enclosed in a large vacuole filled
with a mucilaginous substance. The degree of concentration of
the mother from which the crystals have separated, deter-
liquor
mines, according to KNY ( ), their form, whether tetragonal or
71

monoclinic.

SILICEOUS BODIES, which are only soluble in hydrofluoric acid, are found in the
cytoplasm of many cells, especially of Palms and Orchids, and often completely
(ill the whole cell.

TANNIN. Highly refractive vacuoles filled with a concentrated solution of

tannin are of frequent occurrence in the cytoplasm of cortical cells, and may often
grow to a considerable size. The dark-blue or green colour reaction obtained on
treatment with a solution of ferric chloride or ferric sulphate, and the reddish-
brown precipitate formed with an aqueous solution of potassium bichromate, an:
usually accepted as tests for the recognition of tannin, although equally applicable
for a whole group of similar substances.
FATS and OILS in plants are mixtures of fatty-acid esters. Frequently, a> in
most Monocotyledons, a fatty oil appears in the old chlorophyll grains. The
occurrence of castor in the form of highly refractive drops in the cytoplasm of
oil

the aleurone-containing cells in the endosperm of the castor-oil seeds, has already
been referred to. Oil usually occurs in this form. But fatty substances may also
appear in the cytoplasm as irregularly shaped, more or less soft grains, as for
example in the vegetable butters and in the wax of various seeds they may even;

be crystalline, as in the needle-like crystals of Para-nuts (Bertholletid exwlsa) and of

Nutmeg (Myristica fragrans). Special cytoplasmic structures in which the drops
ofoil are embedded, are of frequent occurrence in the epidermal cells of Oreliidac < -.n-

and Liliaceae and go by the name of elaioplasts 72 ). (

CHOI.KSTK.UIN. Since this substance is a regular constituent especially of givi-n

probably plays an important part in metabolism ( ).
7:i
cells, it
(ii.vccici.N. This substance ( 74 ), related to starch, and of frequent occurrence
in animal tissues, fulfils, according to EKKKKA ( 75 ), the same functions in the

Fungi as sugar and starch in the higher plants. Cytoplasm containing glycogen
iscoloured a reddish brown with a solution of iodine. This colour almost wholly
disappears if the preparation be warmed, but reappears on cooling.
 MORPHOLOGY 79

ETHEREAL OILS AND RESINS. In many cases the strongly refractive drops
found dispersed throughout the cytoplasm are globules of some ethereal oil. It is
the presence of such oils in the petals of many flowers that gives to them their
agreeable perfume. Under certain conditions the oil globules may become crystal-
lised. This occurs, for example, in Rose petals. In 7iio.st cases ethereal oils or
resins are formed in special protuberances ( 76 ) or strata ( 77 ) of the cell wall and only
later are stored in the interior of the cell or in special intercellular spaces formed

hy separation or destruction of cells. Special cells of this kind, often with corky
walls and filled with resin or ethereal oils, are found in the rhizomes of certain
plants, as for instance -in those of Acorus Calamus and of Ginger (Zingibcr
qfficinalc) : Cinnamon trees (Cinna'niomum) ;
also in the bark, as, for example, of
iu the leaves, as in theSweet Bay (Laurus nolnlis) in the pericarp and seed of
;

the Pepper (Piper niyrum) in the pericarp of Anise (Illicium anisatwn).

;

Mn ILACJK is often found in the cells of bulbs, as in Alt turn Cepa and L'njiimi
Xcilla, in the tubers of Orchids, also in aerial organs,especially in the leaves of
succulents, which, living in dry places, are thus enabled to maintain their water
supply by means of their mucilaginous cells.
CAOUTCHOUC AND GUTTA-PKRCHA. These substances are found in a number of
plants belonging to different groups, in particular in the Moraceae, Euphorbiaceae,
and Sapotaceae. They occur in the latex of special cells in the form of small
globules, which, suspended in
the watery sap, give it its milky appearance.
FERMENTS. Bodies of this nature are widely spread in vegetable cells. Their
significance in the oxidation processes taking place in the organism
has been dealt
with by R. CHODAT, A. BACH (
78
), and others. The name leptomin was given
by RACIHOUSKI
9
(~ ) to a catalytic enzyme,
which he found in the sieve-tubes and
laticiferous elements of the higher plants, in the milk of the coco-nut, and in|the
tissues of the potato tuber.
SULPHUR. The presence of sulphur in the form of small refractive grains in
the protoplasm of certain Bacteria, the Beggiatoae, is noteworthy. These Bacteria
80
live in water containing much organic matter, and, according to WINOGRAPSKY ), I

obtain their sulphur from sulphuretted hydrogen. In fulfilling its function in the
Bacteria the sulphur becomes oxidised into sulphuric acid.

The Cell Sap. Under this term is included especially the fluid
which the inner sap cavity.
in old cells fills It is generally more
and clearer than the fluid contained in the smaller vacuoles
watery
of the cytoplasm. No sharp distinction can, however, be drawn
between the sap cavity and vacuoles, and, moreover, a number of
such vacuoles take the place of the sap cavity itself.
may The cell
an acid reaction, owing to the presence in it of
sap usually gives
organic acids or their salts. The substances held in solution by the
cell sap are very various. The soluble carbohydrates, in particular
the sugars (cane sugar, the glucoses, and especially grape sugar),
frequently occur in the cell sap. The glucoses may be recognised by
their reducing properties.

If preparations containing glucose be placed in a solution of copper sulphate,

and, after being washed out, are transferred to a solution of caustic potash and
heated to boiling, they will give a brick-red precipitate of cuprous oxide. If cane

.sugar or saccharose be present, this same

treatment gives only a blue colour to the
cell sap.
80 BOTANY r.urr i

Carbohydrates are transported in a plant principally in the form

of glucose ; cane sugar, on the contrary, is stored up as a reserve
material, as for example, in the sugar-beet, in the stems of sugar-
cane, and in other plants from which the sugar of economic use is
derived.

INULIN', a carbohydrate in solution in cell sap, behaves in the same way in the
(
''ompositae. Treated with alcohol, inulin is precipitated in the form of small granules,
which may be redissolved in hot water. When portions of plants containing much
inulin, such as the root tubers of Dahlia variabilis, are placed in alcohol or dilute
glycerine, the inulin crystallises out and forms sphserites, spheroidal bodies com-
posed of radiating crystal needles arranged in concentric layers. Amides such as
GLUTAMIN and ASPARAGIN are also generally present in the cell sap. There are
frequently found dissolved in the cell sap TANNINS, ALKALOIDS, and GLUCOSIDES.
such as coniferin, hesperidin, amygdalin, solanin, tvsculin, saponin, and also bitter
principles related to the glucosides. Organic acids (malic, formic, acetic, and
oxalic acids) are also of frequent occurrence in the cell sap thus, malic acid is
;

usually present in the leaves of the succulents. For the most part, these organic
acids unite with bases, and the salts which are formed often crystallise. Of acid
salts, which are than free acids, the binoxalate of potassium found in
less frequent
Field Sorrel (Rumex) and Wood Sorrel (Oxalis) deserves special mention. Species
of Salicornia and Salsola contain sodium oxalate. The cell sap always contains
dissolved inorganic salts, especially nitrates, sulphates, and phosphates. The
different vacuoles of the same protoplast may have distinct contents thus one
;

may contain tannin and another be free from it, or one may have coloured and
another colourless sap.

The cell sap is often coloured, principally by the so-called ANTHO-

CYANIN, which Hans Molisch, at least in some cases, regards as being
a non-nitrogenous glucoside ( 81 ). This is blue in an alkaline, and red
in an acid reacting cell sap, and, under certain conditions, also dark

red, violet, dark blue, and even blackish-blue. Anthocyanin can be ob-
tained from the super-saturated cell sap of a number of deeply coloured
parts of plants in a crystalline or amorphous form. Blood-coloured
leaves, such as those of the Copper Beech, owe their characteristic
appearance to the united presence of green chlorophyll and antho-
cyanin. The different colours of flowers are due to the varying
colour of the cell sap, to the different distribution of the cells
containing the coloured cell sap, and also to the different com-
binations of dissolved colouring matter with the yellow, orange,
or red chromoplasts and the green chloroplasts. There is occa-
sionally found in the cell sap a yellow colouring mattter known
as xanthein ;
it is nearly related to xanthophyll, but soluble in
water.

2. ONTOGENY OF THE CELL

The Origin of the Living Elements of Protoplasm. All the
nucleiin an organism owe their origin to the nuclei of previous
SECT, i MORPHOLOGY 81

generations. The spontaneous formation of a nucleus never takes

place. In the same manner, the cytoplasm of every organism is
derived from the cytoplasm of the germ cell, and, so far as is yet
known, the chromatophores take their origin only from their own
kind.
Nuclear Division. Except in a few limited cases, nuclei reproduce
themselves by MITOTIC or INDIRECT DIVISION. This process, often
referred to as KARYOKINESIS, is somewhat complicated, but seems
necessary in order to effect an equal division of the substance of the
mother nucleus between the two new daughter nuclei.
Indirect Nuclear Division ( 82 ). In its principal features the pro-
cess is similar in the more highly organised plants and in animals.
Its stages are represented in a somewhat diagrammatic manner in the

following figure (Fig. 86), as they occur in a vegetative cell such

as those which compose the growing point.
The fine network of the resting nucleus (Fig. 86, 1 n) becomes
drawn together at definite points and separated into a number of
bodies (Fig. 86, 2 ch), the outline of which is at first irregular. Their
form soon becomes filamentous, and the filaments become denser and
at the same time shorter and thicker (3), and stain more deeply.
The stainable substance of the filament, which is called chromatin,
becomes arranged in more or less regular transverse discs united by
linin (3). The filaments themselves are called CHROMOSOMES (Fig. 86,
3, 4). The chromosomes are moved into the plane of division where
they constitute the nuclear or equatorial plate (5 kp, 6, 7). Each
chromosome has meanwhile undergone a longitudinal split which
continues to become more marked (5, 6, 7). The two halves of
each chromosome thus separated move away from one another in
opposite directions, and take part in the formation of the daughter
nuclei (9 t).
Other changes serve to direct the process thus briefly described.
While the nuclear network is separating into the individual chromo-
somes, cytoplasmic filaments become applied to the nuclear membrane,
surrounding it with a fibrous layer. This layer becomes raised up
from the nuclear membrane at two opposite points (3 k) and forms the
polar caps. These are filled with a homogeneous substance in which
fine filaments appear later. The latter converge at the poles, without,
however, coming into contact ; they constitute two pointed bundles,
since they diverge from one another as they pass from the polar
regions (4 &). At this stage the nucleoli (nl) are dissolved and the
nuclear membrane disappears. The fibres proceeding from the polar
caps can thus become prolonged into the nuclear cavity (4, 5). Here
they either become attached to the chromosomes, or^ filaments from
the two poles may come into contact and extend continuously from
the one pole to the other. In this way the nuclear spindle is formed
(5, 6, 7). The fibres of the spindle attached to the chromosomes may
 BOTANY

be termed traction-fibres, those which run from pole to pole supporting-

fibres. The nucleoli appear to form a reserve substance which serves
to nourish the chromosomes and, later, has the special duty of provid-
ing material for the formation of the spindle. Any excess of nucleolar
substance passes into the surrounding cytoplasm, where it forms the

FIG. 86. Successive stages of nuclear- and cell-division in a meristematic cell. Nucleus; nl,
.,

nucleolus ; tc, nuclear membrane; e, cytoplasm; eh, chromosomes; k, polar cap-.; s, spindle;
kj>, nuclear plate t, young daughter nuclei
; c, connecting fibres : z, cell-plate ; m, new cell
;

wall. In 1, the resting nucleus 2 and 3, separation of the chromosomes ; 4, chromosomes with
;

transverse discs 5, the arrangement of the chromosomes to form the cell plate and their
;

longitudinal fission ; 3-5 show the formation of the spindle from the polar caps r., the
;
longi-
tudinal fission of the chromosomes 7, the beginning of their separation to either pole 8, the
; ;

complete separation of the daughter chromosomes; 9, jwssage of the daughter chromosomes

to either pole 10-12, formation of the daughter nuclei in 9-11 the origin of the connecting
; ;

fibres and of the cell plate is seen, while in 12 the new cell-wall is formed. ( x alnnit 000.)

so-called extranuclear nucleoli. The traction -fibres promote the

arrangement of the chromosomes in the equatorial plane (5). After
the chromosomes have split longitudinally into the daughter chromo-
somes, the latter are separated and drawn towards the two poles by
the contraction of the traction-fibres (8, 9). The supporting-fibres
afford the necessary resistance in the process. The spindle fibres can
often be traced to the limiting layer of the cytoplasm, and their
 MORPHOLOGY 83

attachment to this determined. In forming the daughter nuclei, the

free ends of the chromosomes become drawn in (10), and the
first

surrounding cytoplasm separates itself by means of a protoplasmic

membrane, the nuclear membrane (11), from the developing nuclei.
Within the nuclear cavities which are thus produced, the chromo-
somes again assume a reticulate structure and unite with one another

KII:. S 7. Diagrammatic representation of the different positions occupied by the chromosomes iu

the spindle and daring their separation, a and b, daughter chromosomes of one mother
chromosome ; z, traction-fibres of the spindle.

to form a network within which their individual limits are not dis-
tinguishable. We are compelled, however, to assume that the
individuality of the chromosomes is not lost ( 83 ). The young nuclei
enlarge, the extranuclear nucleoli disappear in the surrounding
cytoplasm, and one or more nucleoli at length appear in the nuclei ;
finally the resting condition is again attained.

The process of nuclear division is described above as it usually takes place iu

the young tissues of more highly organised plants. The chromosomes are usually
loop-shaped, and the traction-fibres become attached to the middle point of the
loop, less commonly toward an end. The traction-fibres from the two poles attach
themselves respectively to the two daughter chromosomes of each pair. The
orientation of the pairs of daughter chromosomes iu the nuclear spindle, and the
way in which they separate from one another, are represented in the accompany-,
ing diagram (Fig. 87). In (1) the two shanks of each pair of chromosomes lie
nearly in the equatorial plane, and each of the daughter chromosomes on their
separation (2) assumes the form of the letter U, remaining attached by the shanks.
More commonly, while one of the shanks of the paired chromosome lies in the
equatorial plane, the other is directed towards one of the two poles (3). In this
case a condition of things results, when the daughter chromosomes separate, which
is represented in 4 or 5. The former shows the resulting appearance when the
daughter chromosomes remain for a time attached to one another at both ends ;

the latter separation soon follows at the end directed towards the nuclear
when
pole. In all cases the separation proceeds from the point of attachment of the
traction-fibres. When a paired chromosome is attached to the spindle near one of
its chromosomes naturally commences near
ends, the separation of the daughter
thisend when the attachment is by the middle of the chromosome the daughter
;

chromosomes remain longer attached by their ends. In Fig. 86 the behaviour of

84 BOTANY PAKT i

the chromosomes is represented as in the diagrams 3 and 4 (Fig. 87). As a rule it

does not appear so clearly, but more or less combined with the other type.
The changes occurring in a mother nucleus preparatory to division are termed
the PROPHASES of the karyokinesis. These changes extend to the formation of the
nuclear plate, and include also the process of the longitudinal division of the chromo-
somes. The stage of the nuclear plate is the METAPHASK. The separation of the
daughter chromosomes is accomplished in the ANAPHA.SE, and the formation of
the daughter nuclei in the TELOPHASE of the division. The real purpose of the
whole process is attained in the quantitative and qualitative division of the
chromosomes, resulting from their longitudinal splitting (Fig. 86, 5, 6, 7 ;

Fig. 87). The anaphases and telophases of the karyokinesis are but a reverse
repetition of the prophases.
The number of chromosomes occurring in any nucleus is a definite one, and \vla-n
a deviation from the usual number is met with, it is due to some of the chromo-
somes having remained united end to end. The smallest number of chromosome
which has yet been found in the nuclei of vegetative cells of the more highly
organised plants has been eight as a rule the number is larger, amounting often
;

to several times this number.

A special type of nuclear division, to which the name of REDUCTION

DIVISION is
given,, is met with in those reproductive cells which start a
new generation, such as the spore-mother-cells of the higher Cryptogams
and Phanerogams. In the prophase of this division the chromosomes
become united in pairs (Fig. 88, 1, 2), and there then occurs a mai'ked
contraction of the nuclear contents, which is characteristic of this
process of division and is called SYNAPSIS (3). After this the double
chromosomes become again loosened out as a delicate double thread
(4), which soon unites to
a correspondingly stout thread, forming a
loose skein (5). The doubled nature of this thread soon becomes
recognisable again (6). The skein consisting of the as yet unbroken
double thread now falls into segments (7), each of which corresponds
to one paired chromosome. The number of these segments is half
as great as the number of chromosomes in the tissue cells of the
same plant, since two chromosomes are represented by each segment.
The paired chromosomes become shorter and thicker and are
distributed around the periphery of the nucleus this is the con-
;

dition that has been termed diakinesis (8). At this stage kinoplasmic
filaments are becoming applied to the nuclear membrane (8) ; the
latter disappears and the nuclear spindle, which is at first multipolar

(9), but ultimately becomes bipolar (10), originates from the kino-
plasmic fibres. The paired chromosomes become attached to the
fibres of the spindle and arranged in an equatorial nuclear plate (10).

Shortly afterwards the separation of the chromosomes, until now

united in pairs, takes place (11). In this process, in which the
essential of the reduction division is effected, it is not longitudinal
halves of chromosomes but entire chromosomes which separate from
one another. The result of this is that each daughter nucleus receives
only half as many chromosomes as were found in the tissue cells of the
 MORPHOLOGY 85

IS

;. S8. Dividing polleii-iiiother-cells of a Lily, somewhat diagrammatic. 1, Mother-cell with

resting nucleus ; 2, the separation of the chromosomes 3, synapsis ; 4, double filament in process
;

of fusion ; 5, spirem consisting of an apparently single filament derived from the fused double
filament 6, reappearance of the longitudinal split, the spirem still unsegmented 7, spirem
; ;

transversely segmented, into paired chromosomes ; 8, diakinesis 9, multipolar spindle

; 10, ;

spindle of the mother-nucleus, the nuclear plate composed of paired chromosomes ; 11, re-
duction division, the separating chromosomes showing partial separation of their longitudinal
halves ; 12, young daughter nuclei 13, the longitudinal halves of the chromosomes (daughter
;

chromosomes) are arranged in pairs on the nuclear spindles 14, the spindles of the daughter
;

nuclei ; 15, separation of the daughter chromosomes ; 16, young stage of the grand-daughter
nuclei, (x about SOO.)
 86 BOTAXY

same plant. During their passage towards the poles of the spindle
a longitudinal split can be detected in each chromosome. This split
was indeed already complete in the prophase before the nuclear plate
was formed, but was not followed as in an ordinary division by a
separation of the halves. The two halves of each chromosome
remain on the other hand in relation to one another and pass to the
? same daughter nucleus. The formation of the daughter nuclei is
completed (12) as in an ordinary division, but following promptly on
the first reduction division, which is also known as the HETEROTYPE
84
division, comes a second or HOMOTYPE division ( ). In this no new
longitudinal splitting of the chromosomes takes place, but the two halves
of each chromosome, which existed in the daughter nuclei, become
separated from one another, and become the chromosomes of the
grand-daughter nuclei.
The steps of this homotype division agree in other respects with
those of an ordinary nuclear division, and will be clear from Fig. 88,
13-16. In 13 an early 'stage and in 14 the completed condition of
the spindles of the dividing daughter nuclei are seen; 15 shows the
division of the nuclear plate, and in 16 the young grand-daughter
nuclei are completed. One of the characteristic features of the whole
is that the two divisions succeed one another immediately
process
or very quickly. The heterotype and homotype nuclear divisions,
which may together be termed the ALLOTYPIC division, may be con-
trasted with the ordinary or typical nuclear division. At a particular
stage of development corresponding phenomena to those of the allo-
typic division are met with in animals as well as plants.
The smallest reduced number of chromosomes known for the nuclei of the
more highly organised plants is four, i.e. the
half of the smallest number met with in the
tissue cells.
In those lower Cryptogams, the nuclei of
which possess an individualised centriole, the
latter undergoes division into two at the be-
,"> v. j j; -

* .-
'

, -.'>," ,
ginning of karyokinesis. The two halves
*
^.y: '..-.; pte from one another (Fig. 89 c) and
X-^ltfePRSr^? ultimately reach the jioints which will become
ultimate!
^rV^ESag ~'
-

the pole,
poles of the spindle. Round such cen-
trioles a definite portion of
protoplasm form-
^LyT/'-v ;
?~\ ing the centrosome is usually marked off,
and around this kinoplasmic radiations (kp)
FIG. MI. A nucleus of a vouii" plant of v j_- i
the Brown Seaweed, Fitcus serratus,
,.
form an astrosphere. When j.i
IITI.
the centnoles

preparing to divide. The two centrioles

have reached the poles the nuclear membrane
(c), which have arisen l.y the division disappears, and spindle fibres appear in the
of a sin-le one, have already separated nuclear These clearly proceed
cavity itself.
from one another;/.-?, radiations of the
from the cent rosomes and become atta.-h.-<i
nbnllar plasma; s, chromosomes
to
;

e chromosomes. ^ _,
The complete nuclear 1.1.1
nucleolus (x 1000)
spindle (Fig. 90) has a centrosome with kino-
lasmatic radiations (kp] at each pole, but in other respects agrees with the spindles
 MORPHOLOGY 87

of the higher plants, from which centrioles are absent. The main features of
the division and the formation of the daughter nuclei are also similar in the two
cases. The centrioles persist in the daughter nuclei
and .divide into two on each subsequent nuclear
division ;
the kinoplasmatic radiation around the
-kj,
centrosome is, however, only present during the -C
M ).
karyokinetic process (

Direct Nuclear Division. In addition

to the mitotic or indirect nuclear division
there is also a DIRECT or AMITOTIC division,

sometimes called FRAGMENTATION (Fig. 91).

This may have been the original mode of
nuclear division, and among the lowest
organisms transitions between it and in-
direct division are found. In the higher
FIG. !K). A nuclear spindle of a
plants direct division of the nucleus is a cell of a young plant of the
reduced, or indeed a senile process which Brown Seaweed Fm-us serrd.tvs
with split chromosomes in tin-
usually occurs in old cells, or in cells in nuclear plate, c, Centrioles
which the contents become disorganised*
cell /,
/i, radiations of fibrillar plasma

shortly after the nuclear division. astrospheres; sp, spindle

fibres;longitudinally divided
Instructive examples of direct nuclear 8,
chromosomes funning the
division are afforded by the long internodal nuclear plate, (x 1000.)

cells of the Stoneworts (Characeae), and also

by the old internodal cells of Trades-
cantia (Fig. 91). The direct nuclear divi-
sion is chiefly a process of constriction
which, however, need not result in new
nuclei of equal size. In the case of the
Stoneworts, after a remarkable increase
in the size of the nucleus, several
successive rapid divisions take place, so
that a continuous row of bead-like nuclei
results. The old internodal cells of
Tradescantia (Fig. 91) very frequently
show half-constricted nuclei of irregular
shape. While in uninuclear cells indirect
nuclear division is, as a rule, followed
by cell division, this is not the case after
direct nuclear division.
Cell Division. In the uninuclear
cells of the higher plants cell division
FKI. 9i. -old cells from the stem of and nuclear division are, generally, closely
virginica, showing associated. The supporting fibres of
nuclennpro, ,

pole persist as CONNECTING FIBRES

between the developing daughter nuclei (Fig. 86, 90; Fig. 88, 12, 16),
 BOTANY

and number is increased by the interposition of others (Fig.

their
86, 10). In consequence of this a barrel-shaped figure is formed,
which eithe/ separates entirely from the developing daughter nuclei,
or remains in connection with them by means of a peripheral sheath,
the CONNECTING UTRICLE. The first is the case in cells rich in cyto-
plasm, the latter when the cells are more abundantly supplied with
cell sap. At the same time the con-
necting fibres become thickened (Fig.
86, 10) at the equatorial plane, and
the short rod-shaped thickenings form
what is known as the CELL PLATE.
In the case of cells rich in proto-

plasm or small in diameter, the con-

necting fibres become more and more
extended, and touch the cell wall at
all points of the equatorial plane (Fig.
Fio. 92. Three stages in the division of a
86, 11). The elements of the cell
living cell of Epipactis paliistrls. (After
TREUB, x 365.) plate unite and form a cytoplasmic
limiting layer, which then splits into
two. In the plane of separation the new partition wall is formed
of cell-wall substance, and thus SIMULTANEOUSLY divides the mother
cell into two daughter cells (Fig. 86, 12). If, however, the mother cell
has a large sap cavity, the connecting utricle cannot at once become so
extended, and the partition wall is then formed SUCCESSIVELY (Fig.
92). In that case, the partition wall first commences to form at the
point where the utricle is in contact with the side walls of the mother
cell (Fig. 92 A}. The protoplasm then detaches itself from the part
of the new wall in contact with the wall of the mother cell, and
moves gradually across until the septum is completed (Fig. 92 B
and 0) ; the new wall is thus built up by successive additions from
the protoplasm. *

The connecting fibres of the spindle consist of kinoplasm. The origin of the
cell platefrom swellings of these fibres indicates its kinoplasmatic nature also. By
the splitting of the cell plate the limiting layers of the two sister cells are com-
pleted across the place of separation. The separation of the complex of connecting
fibres into two halves effects an equal division of the kinoplasm between the two
new cells.

In the Thallophytes, even in the case of uninuclear cells, the

partition wall is not formed within connecting fibres, but arises
either simultaneously from a previously formed cytoplasmic plate, or
successively, by gradual projection inwards from the wall of the
mother cell. It was a division process of this kind (Figs. 93, 94),
first investigated in fresh-water
Algae, that gave rise to the conception
of cell division, which for a long time prevailed in both animal and

vegetable histology. In this form of cell division the new wall com-
SECT. I 89

mences as a ring-like projection from the inside of the wall of the

mother cell, and gradually pushing farther into the cell, finally extends

FIG. 93. Cell of Spirogyra in division. FIG. 94. Portion of a dividing cell of Clado-
n, One of the daughter nuclei w, de-
; phora fracta. w, Newly-forming partition
veloping partition wall ch, chloroplast
; wall ; eh, dividing chromatophore ; k,
pushed inward by the newly-forming nuclei, (x 600.)
wall. (X 230.)

completely across it (Figs. 93, 94). In a division of this sort, in

uninuclear cells, nuclear division precedes cell division, and the new
wall is formed midway be-
tween the daughter nuclei
(Fig. 93). In the multinuclear
cells of the Thallophytes, on
the other hand, although the
nuclear division does not differ
from that of uninuclear cells,
cell division (Fig. 94) is al-

together independent of nuc-

lear division. And in multi-
nuclear, unicellular Thallo-
phytes, nuclear division is not
followed by a cell division.
The interdependence of
nuclear and cell division in
uninuclear cells is necessary
to ensure a nucleus to each >

daughter cell. In multi- -SffillS

nuclear cells it is not essen-
tial that cell division should

always be accompanied by
nuclear division, as in any FIG. 95. Portion of the peripheral protoplasm of the
Case a Sufficient number of embryo-sac of Keseda odorata, showing the commence-
T .,,. ,. , , ment of multicellular formation. The process pro-
nuclei will be left to each downwards (x 240 .)
gresses from above .

daughter cell.

Free Nuclear Division and Multicellular Formation. The nuclear division in

90 BOTANY TAUT I

the multinuclear cells of the Thallophytes may serve as an example of free nuclear
division, that is, of nuclear division unaccompanied by cell division. In plants
with typical uninuclear cells, examples'of free nuclear division also occur although. ;

in that case, the nuclear division is customarily followed by cell division. While
the nuclei increase in number by repeated division, this process is not accompanied
by a corresponding cell division. When, however, the number of nuclei is com-
pleted, the cytoplasm between the nuclei divides simultaneously into as many
portions as there are nuclei. In this process we have an example of multicellular
formation. This method of development is especially instructive in the embryo-
sac of Phanerogams, a cell,
often of remarkable H/C
and rapid growth, in which
the future embryo is de-
veloped. The nucleus of
the rapidly growing em-
bryo-sac divides, the two
daughter nuclei again
divide, their successors
repeat the process, and so
on, until at last thousands
of nuclei are often formed.
Xo cell division accom-
panies these repeated nuc-
lear divisions, but the
nuclei lie scattered
throughout the peripheral,
cytoplasmic lining of the
embryo-sac. When the
embryo-sac ceases to en-
large, the nuclei surround
Fio. 96. Successive stages of the delimitation of a spore in the themselves with connect-
ascus of Erysiphe eommunis. A, Before delimitation has ing strands, which then
begun the tibrillar plasma (kp) radiates into the cytoplasm radiate from them in all
;

around; in B the fibrillar plasma has commenced to grow

directions ( Fi g- Cell
round the mass of cytoplasm in C this process is com-
; ^5).
plete, but the limiting layer produced by the fusion of radia- plates make their appear-
tions of the fibrillar plasma is still connected with the polar ance in these connecting
side of the nucleus ; in D this kinoplasniatie connection strands and from them
between nucleus and limiting layer has disappeared n T~ tins
nuclear network ; n, nucleolus.
,'r.HARPER,
(After x 1500.)
; *, 11
cell walls arise.
;
In <-\

manner the peripheral

protoplasm of the embryo-sac divides, simultaneously, into as many cells as there
are nuclei. Various intermediate stages between simultaneous, multibellular forma-
tion and successive cell division can often be observed in an embryo-sac. Win-re
the embryo-sac is small and of slow growth, successive cell division takes place,
so that multicellular formation may be regarded as but a shortened process of
successive cell division, induced by an extremely, rapid increase in the size of
the cell.
Free Cell Formation. Cells produced by this process differ conspicuously from
those formed by the usual mode of cell division, in that the free nuclear division is
followed by the formation of cells, which have no contact with each other, and in
the formation of which the whole of the cytoplasm of the mother cell is not used
up. This process can be seen in the developing embryo of the Gymnosperms, in
 SECT, i MORPHOLOGY 91

for example, and also in the formation of the

Ephedra, spores of the Ascomycetes.
A single nucleus is present to begin with in each ascus of the Ascomycetes. By
successive divisions eight nuclei lying free in the cytoplasm are derived from this.
A definite portion of cytoplasm around each of these nuclei becomes limited by a

peripheral layer which then forms a cell wall. Thus eight separate spores arise (cf.
Fig. 327). As the researches of Harper ( 86 ) have shown, the formation of the peri-
pheral layer proceeds from a centrosome-like mass of kinoplasm which formed a pole
of the spindle in the preceding nuclear division. The nucleus is drawn out towards
this mass of kinoplasm. From the latter kinoplasmic radiations proceed which
surround the spore as it becomes delimited, and finally fuse to form its peripheral
layer (Fig. 96).
Cell - Budding. This is simply a special variety of ordinary cell division, in
which the not divided in the middle, but, instead, pushes out a protuberance
cell is

which, by constriction, becomes separated from the mother cell. This mode of cell
multiplication is characteristic of the Yeast plant (Fig. 2, p. 11) and the spores, ;

known .as conidia, which are produced by numerous Fungi, have a similar origin
(Fig. 353).
Conjugation (
87
). A sexual cell
is, with few exceptions, only able to continue
its development with another sexual cell. The two cells so uniting
after fusion
are either alike, and in that case are called GAMETES, or unlike, and are then dis-
tinguished as EGG and SPERMATOZOID. The spermatozoid is the male, the egg the
female sexual cell. The gametes may be motile (Fig. 97 B) or non-motile. The
motile gametes frequently resemble the swarm-spores (Fig. 97 A) generated by
the same parent for the purpose of asexual reproduction. As a rule, however, they
are smaller than the and have usually only half as many cilia. In
swarm-spores,
the more highly specialised sexual cells the egg usually retains the structure of an
embryonic cell, but the spermatozoid undergoes profound modifications. A
cytoplasmic body, a nucleus, and the rudiments of chromatophores are
cell

always present in the egg. The spermatozoid (Fig. 98), on the other hand,
becomes transformed, in the more extreme cases, into a spirally twisted body,
provided with cilia, and exhibiting an apparently homogeneous structure. Only
a knowledge of the history of its development, and the greatest care in hardening
and staining, have rendered it possible to recognise the homology of the structure
of such a spermatozoid with that of an embryonic cell. It has been shown that
the hinder part of its spiral body corresponds to the cell nucleus (k), the anterior,
together with the cilia, to the cytoplasm, especially the kinoplasm (c), and the
vesicle (I), at the other extremity, to the sap cavity of a cell ( 8tJ ).
Motile male cells provided with cilia occur only in the Cryptogams and, as has
89
been recently demonstrated ( ), in some Gymnosperms (Cycadaceae, Ginkgo). In
the Cryptogams the spermatozoids are set free from the sexual organs and require
water for their dispersal. They reach the egg-cell, which usually remains in its
place of origin, by swimming. In the Gymnosperms, which form motile spermato-
zoids, the latter are brought near to the ovum by means of the pollen tube developed
from the pollen grain. In a similar way the non-motile male cells of the other
Gymnosperms and the Angiosperms are conducted to the egg through the pollen
tube (Fig. 99). In the union of the two sexual cells in the act of fertilisation, the
egg nucleus (ek) and the sperm nucleus (sk) fuse and form the nucleus of the fertilised
egg-cell. The cytoplasm of the male cell also commingles with that of the female
cell, but the chromatophores of the embryo are derived from the egg-cell alone.
When the spermatozoid, as in animals and in Thallophytes, is provided with a
centrosome, this does not fuse with the centrosome belonging to the ovum. The
92 BOTANY

centrosome of the fertilised egg- cell appears to be derived from that of the
.s[>ermatozoid only.

FIG. 97. A, An asexual swarm-spore

of Ulothrix zonata ; B, \, a gamete ;
2 and 3, conjugating gametes ; 4,

zygote, formed by the fusion of

two gametes, (x 500.)

Kio. 00. Fertilisation of a phanerogamic Angio-

sperm, somewhat diagrammatic. .-1, End of
FIG. 98. A, Spermatozoid of Cham pollen tube in it the generative cells gz, each
;

fruijilis ; B, Spennatozoid of the of which contains a sperm nucleus ://., til--

Fern, Onodeastruthiopteris. (After vegetative nucleus in process of dissolution.

SHAW.) The nucleus (k) is more B-D, Egg in successive stages of fertilisation,
deeply shaded ; c, the cytoplasmic B, showing the generative cell with its
portion ; c7, cilia which in B arise sperm nucleus, nk, penetrating the egg; syn,
from the tinner edge of the spiral the degenerating synergidae; C, the union of
band ; 6, vesicle. (A x 540, sperm nucleus, sk, and egg nucleus, ek ; D,
JB X 850.) the germ nucleus, kk, resulting from the
fusion of the sperm and egg nuclei; cJi,
rudiments of chromatophorcs. ( x about 500.)

The nuclei of the sexual cells always possess only the reduced number of
chromosomes. By fertilisation the number becomes doubled, and after a shorter
or longer period is again halved in the reduction division.
 MORPHOLOGY 93

Although it is by fertilisation that the further development of the sexual

products is made possible, there are exceptional cases in which a gamete or an egg
forms an embryo without being fertilised. If development follows in this way from
a gamete or egg with the reduced number of chromosomes the phenomenon is
calledPARTHENOGENESIS. On the other hand the reduction of the number of
chromosomes may be suppressed in the development of the egg so that the full
number necessary for further development is already present in it. In this case
its development is not true parthenogenesis but a vegetative production of an
embryo, which has come about through loss of sexual-
ity and falls under the conception of APOGAMY ( ).
go

The cases of production of embryos from unfertilised

eggsamong the higher plants belong to the latter category :

this isknown in Antennaria alpina ( 91 ), Alchemilla ( 92 ),

Thalictrum purpurascens 93 ), Taraxncum w ), Hieracia ( 95 ).
( (

In lower plants, in some Fungi, e.g. Saprolegnieae, and in

one of the Characeae (Chara crinita) ( M ) true partheno-
genesis is found. In Marsilia ( 97 ), one of the Hydro- FJG 10o.
Chlorophyll
pterideae, the segmentation and further development of grains from the leaf of
unfertilised eggs has also been observed. Funaria hygrmnetriva,
resting, and in process
Multiplication of the Chromatophores. This of division, smaii m-
eluded stan* grains are
is accomplished
r by a direct division, as a result
. '. present in the grains.
of which, by a process of constriction, a chro- ( X 540.)

matophore becomes divided into nearly equal

halves. The stages of this division may best be observed in the
chloroplasts (Fig. 100).

B. Cell Fusions

The connection of the living protoplasts with one another is less

complete in plants than in the animal body. This is evident from the
existence of the cell wall which surrounds the vegetable protoplast.
Recent researches have, however, shown ( 9S ) that the protoplasts of
the plant are united together by extremely fine cytoplasmic filaments,
which proceed from the boundary layer of the cytoplasm. Such
filaments are mostly confined to the pit-membrane (Fig. 101), but
may also penetrate the whole thickness of the cell wall (Fig. 102).
The existence of these connecting filaments of living substance
between the protoplasts confers an organic unity on the whole body
of the plant.
The members which make up the sieve-vessels, or, as they are
commonly called, the sieve-tubes, are united by thicker strands of
cytoplasm, which facilitate the transfer of substances through the
tube. The transverse walls traversed by these strands of cytoplasm
have been referred to above (p. 67) as sieve-plates. The pores
attain their greatest diameter in some Angiosperms (Fig. 103). It
is worthy of special note that, despite the fact that the nuclei of

the sieve-tube segments disintegrate, the cytoplasm, which lines the

94 BOTANY I'AllT 1

wall, remains alive. The walls of sieve-tubes are always unliguitied.

Their sap-cavities contain a watery, and more or less dilute, solution
of albuminous substances, and small starch grains are also present
in most cases. As a rule, the sieve-tubes remain functional only
through one or a few vegetative periods. Before their activity
ceases the sieve-plates become covered (Fig. 103 C) with highly
refractive callus (p. 69).
A more complete fusion of protoplasts is found in the formation
of laticiferous vessels. These have the same structure and contents

, - .

i
pi
'

B
Flo. 10:.'. A, A swollen portion of cell wall
from tin- endosperm ol tlie Vegetable Ivory
-
Palm (1'hyteli ill/a* //m.-,v,r,./y><i). At t, s,

simple pits tilled with cytoplasm ; the

in
~"
intervening pit-membrane are fine connect-
ing threads pi, other threads traversing
;

FIG. 101. A cell from the cortex of the the whole thickness ol the wall, (x 375.)
Mistletoe (Viseum album) the protoplast
; li, The contents of two opposed pits and
has been properly fixed and stained and the connecting threads of the pit -membrane.
the wall (m) swollen. The pit-membranes (X 1500). C, The opening of a pit ami tin-
(x) are traversed by connecting threads ; connecting threads ot the pit-membrane
ck, chloroplasts ; n, nucleus, (x 1000.) viewed from the surface, (x 1500.)

as latex cells (p. 72). Their occurrence, like that of latex cells, is
limited to a few distinct plant families, such as the Papaveraceae, of
which the Poppy (Paparer) or Celandine (Chelidonium), with its
characteristic orange-coloured "sap," are familiar examples, or the

Composited, of which in particular the Lettuce (Lactuca) may be

cited. Latex vessels are distinguished from latex cells only by the
method of their development, which has resulted from the fusion of
rows of elongated cells, the separating transverse walls of which
have become more or less completely absorbed. Such vessels as a
rule possess lateral branches, which, on meeting with one another,
fuse by the absorption of their end walls (Fig. 104). The
61
mucilage tubes ( ) which are found in many Monocotyledons
present points of resemblance to the laticiferous vessels. Their
 MORPHOLOGY 95

mucilaginous sap contains albumen, starch, glucose, and tannins in

addition to inorganic compounds.
A cell fusion also takes place in the formation of VESSELS or
TRACHEA, but it should not be considered as a union between living
cell bodies, but merely as one between cell cavities. The mature
vessels are dead tubes serving for water conduction. They are
formed from rows of cells, the lateral walls of which are peculiarly
marked by spiral or reticulate thickenings, or, as is more frequently
the case, by bordered pits, while the transverse walls become more or
less completely absorbed. In cases where the transverse walls are

FIG. 103. Parts of sieve-tubes of Cucurbita Ptpo, hardened in alcohol. A, Surface view of a sieve-
plate JJ, C, longitudinal sections, showing segments of sieve-tubes
; ; D, contents of two sieve-
tube segments, after treatment with sulphuric acid ; s, companion cells ; it, albuminous con-
tents pr, peripheral cytoplasm <:, callus plate ; c*, small, lateral sieve-pit, with callus plate
:
;

( X 540.)

at right angles to the side walls, they usually become perforated by

a single large round opening while the remains of the wall forms a
thickening ring (Fig. 73 C). When the transverse walls are oblique,
they are perforated by several openings, between which portions
of the wall remain, like rungs of a ladder (Fig. 105 q). According
to the mode of their wall thickening, vessels are distinguished as
SPIRAL, RETICULATE, or PITTED. When the transversely elongated
pits of a vessel are arranged in more or less parallel rows (Fig. 105),
it is called a SCALARIFORM VESSEL. The thickening of the vessel
walls is always lignified. The living contents of the cells, after the
perforation of the transverse walls, become completely absorbed, and
the fully formed vessels or tracheae contain only water and a limited
amount of air.
96 BOTANY

There is no difference between vasit'orm tracheides and vessels other than that
the former are single elongated cells, and the latter fused cell rows. Generally
speaking, tracheides are formed in parts of plants still in process of elongation,
vessels in parts where growth in length has already ceased. True vessels make
their appearance in some of the Ferns, for instance, in the common Bracken
first

(Pteris aquilina). In the main, despite the name Vascular Cryptogams, Ferns
have only vasiform tracheids. Even in the Gymuosperms the Gnetaceae are the
only family regularly provided with vessels. It is in the Angiosperms that vessels
first become of frequent occurrence. Vessels are not of an unlimited length. A
few plants, however, such as the Oak, and especially climbing woody plants, or

Fi<;. 105. Lower third of a scalari-

Km. 104. Tangential section through form vessel from the rhizome
the periphery of the stem of of the common Bracken KITH.
Scorzonera hispanica, showing re- J'ttrif miiiiii mi. Transversely
t,

ticulately united latex vessels. elongated pits in the lateral

(x 240.) walls ; q, scalari form perforations
of the terminal wall. (After DK
BABY, x >.>'.)

Lianes, have vessels several metres long but, as a rule, their length is not more
;

than a metre, and in plants the woody portion of which conducts \\ater only by
vessels, the vessels have an average length of only ten centimetres. The length of
an individual vessel is denned by the presence of transverse walls, which are not
perforated but bear bordered pits.

A similar fusion to that seen above to occur in the formation of

laticiferous vessels is also met with in fungal hyphse by an absorp-;

tion of a part of the wall where two branches come into contact, their
protoplasmic contents unite. A still more complete fusion is exhibited
by the naked amoebae of a Myxomycete in forming the plasmodium
 MORPHOLOGY 97

(p. 56). The fusion of the sexual cells in the process of fertilisa-
tion possesses special characteristics which place the process in a
distinct category.

Tissues (")

Origin and General Properties of Tissues. A continuous

aggregation of cells in intimate union is called a tissue. The origin
of vegetable tissues is, in general, attribut-
able to cell division. It is only in the

Fungi and Siphoneae that a tissue arises

through the interweaving of tubular cells
or cell filaments (Fig. 106). In such cases,
where the filaments are so closely inter-
woven as to form a compact mass of
cells, the tissue thus formed has the same

appearance as the tissues of higher plants

(Fig. 107).
The mutual interdependence of the cells
of a tissue is manifested both by the con- "
Jfi
junction of their pits (Figs. 66, 70, 71),
and by the general similarity of their wall
thickenings.
The cells in a tissue may either fit no. 100. Longitudinal section of
the stalk of the fructification of
closely together (Fig. 70), leaving no open-
ings or spaces, or so-called INTERCELLULAR
SPACES may be left between the individual cells. Where cell fila-
ments are interwoven into a tissue, the intercellular spaces are repre-
sented by the openings left between the loosely-woven filaments (Fig.
106). In tissues resulting from cell division the intercellular spaces
arise subsequently, as the partition wall between two cells formed by
cell division is originally a simple membrane.

Such a partition wall may ultimately split and so give rise to intercellular
spaces, but this only occurs after it has been thickened. The cause of such splitting
is to be found in the hydrostatic pressure existing within the cells, and their con-

sequent tendency to assume a spherical shape. The formation of intercellular

spaces commences, therefore, at the cell corners, where the primary wall, consisting
of pectinose material, becomes swollen.

The simplest and at the same time most frequent intercellular

spaces are triangular or quadrangular in outline, as seen in cross-
section (Figs. 66 i, '74 i). In cases where special portions of
adjoining cells are in extremely energetic growth, intercellular
chambers and passages, of more or less irregular shape, may be
formed between them. If the growth of adjoining cells is very

unequal, it may lead to a complete separation of their cell walls ;

or the cells, or even a whole system of tissues, may be stretched and

H
 98 BOTANY PART i

torn apart. It is by such a process that hollow stems are formed.

Intercellular spaces arising from a splitting of adjoining cell walls
are termed SCHIZOGENIC those formed by
;

tearing or dissolution of the cells themselves

are called LYSIGENIC INTERCELLULAR spaces.
In many cases intercellular spaces that begin
as schizogenic are increased in size by de-
struction of surrounding cells. Most inter-
cellular spaces contain only air, although in

special instances they may contain water or

excreted products, such as gum, mucilage,
resin, or ethereal oils, and in other still
FIG. io7.-Transverse section of
ra rer
Schizogenic intercellu-
cases latex.
the sclerotium of Claviceps ,
.en j -ii. i 11
filled with air, and
lar spaces are usually
purpurea. (x 300.)
form the ventilating system of the plants,
while the lysigenic spaces often contain either water or secretion
products.
Of the schizogenic intercellular spaces, those filled with ethereal oils or resin, on
account of their frequency, should be particularly noticed. Short cavities and
longer passages, or ducts, containing ethereal oils, are to he found in the stems,
roots, and leaves of numerous plant families. The Umbelliferae are especially rich
in these, and the oil-ducts form the characteristic markings (vitte) on their fruits.
The Conifers are especially characterised by resin-ducts (Fig. 141 A, /t), which,
even during their formation by the separation of the cell walls, seem to lill with
the excretion from the cells. The enlargement of such intercellular spaces is
accompanied by a division of the surrounding cells, the number of which is thus
correspondingly increased. The cells themselves remain thin-walled, and in close
contact laterally, but bulge out somewhat into the ducts. Lysigenic intercellular
spaces, acting as receptacles for secretions, have the appearance of irregular cavities in
the tissue. Where they contain oil or resin, they develop from a group of cells in
which these substances appear in the form of drops. The cell group then becomes
disorganised by the gradual absorption of the cell walls. In this way are formed
the receptacles filled with ethereal oils, in the Orange and Lemon. The formation
of the so-called resin-galls, in the case of coniferous trees, is preceded by the
formation of abnormal tissues, which afterwards become converted into resin.
Such was also the origin of amber, which is the fossil resin of the Amber-fir
(Picea succinifera). The formation of gum in lysigenic gum cavities is due to
the modification of the cell walls, and either normal tissues participate in this
process, as in the case of the gum-arabic of the Acacia, or abnormal tissues are
first developed and then transformed into gum, as, for example, the gum on

Cherry trees. Latex does not occur in lysigenic intercellular spaces.

The separating walls resulting from cell division are simple lamellaj. In
tissues which have arisen by cell division these lamellae are common to the cells

they separate. That part of the partition wall between two cells which stands
out so distinctly in a cross-section does not. consist of the original primary cell
wall alone. It is made up of both the primary wall and the primary thickening

layers. The former is called the MIDDLE LAMELLA (Figs. 66 m, 70 m). In soft
tissues the middle lamella, according to DEVAUX ( 10 ), is composed for the mo>t

part of pectic substances ;

in woody and corky tissues it is also lignified. By
 MORPHOLOGY 99

boiling soft tissues in water, the cells may often be easily isolated through the
consequent swelling and dissolutipn of the middle lamella. In ripe fruit, an
isolation of the cells frequently takes place spontaneously, through the dissolution
of the middle lamella. A lignified middle lamella, on the other hand, seems able
to withstand more effectually the action of oxidising agents. Consequently, it
is possible, by subjecting a section of pine -wood to the action of SCHULZE'S
MACEKATING MIXTURE (potassium chlorate and nitric acid), and subsequently
treating with concentrated sulphuric acid, to remove all secondary and tertiary
thickening layers, so that only the middle lamellae remain as a delicate network.
If the macerating process be continued for a longer time, without the subsequent
treatment with sulphuric acid, the middle lamellae become finally dissolved. The
thickening layer will then be left free from all lignification. SCHULZE'S macerating
method may accordingly be employed to isolate the elements of lignified tissues.
The peculiar relation of the middle lamella towards chemical reagents gave rise
at one time to the presumption of a special intercellular substance which, like a

glue,bound together the cells of a vegetable tissue. The supplementary deposition

of pecticcompounds in the middle lamellae (p. 69) frequently gives rise to the
formation of rod -like protuberances and excrescences, which project into the
intercellular spaces, or these spaces may be filled up by the formation of gussets
(Fig. 70 C, m*).

Kinds of Tissue. The mature tissues of the plant may be divided

into two groups, PARENCHYMA and PROSENCHYMA, between which, how-
ever, no sharp distinction can be made. A typically developed paren-
chymatous tissue is one in which the thin-walled cells are equally
expanded in all directions, and possess a layer of protoplasm lining
the wall, and other cell contents. Typical prosenchymatous tissue,
on the other hand, consists of thick-walled, elongated cells, either in
the form of fibres or spindle-shaped cells, with interlocking, pointed
ends, and with little or no protoplasmic contents. A parenchymatous
tissue, in which the cells are thick-walled and elongated, resembles
prosenchyma, but may be distinguished from it by the absence of
pointed cell terminations, and especially by the greater abundance
of protoplasm. Thin- walled prosenchyma is not, on the other hand,
necessarily lacking in protoplasm, but is characterised by its pointed
and interlocking cells.

An undifferentiated tissue, the cells of which are still capable of

division, is termed embryonic tissue, or MERISTEM. The meristem of
embryonic rudiments and of the growing point is called PROMERISTEM,
and all meristematic tissue which can be shown to have been derived
directly from such promeristem is termed PRIMARY. A primary
meristem may still retain its meristematic character in the midst
of a mature tissue. Fully differentiated tissue is designated
PERMANENT tissue, or somatic tissue, in contrast to meristematic tissue.
At times, permanent tissue may again become capable of division, and
thus give rise to a SECONDARY MERISTEM.
A mass of tissue so united in the body of a plant as to form
a higher histological unit constitutes a tissue system. In the more
100 BOTANY PART i

highly organised plants three such systems may be distinguished the

TEGUMENTARY SYSTEM, the VASCULAR BUNDLE SYSTEM, and the
FUNDAMENTAL TISSUE SYSTEM.
The tissues which make up the different tissue systems are
distinguished as PRIMARY and
SECONDARY, according to their
origin. The primary are those which are derived from the pro-
meristem and primary meristems before growth in length has been
completed the secondary are those which owe their origin to the
;

primary or secondary meristems after growth in length is finished.

The primary constituents of the tissue systems will be considered
first.

A. The Primary Tissues

The Tegumentary System. In the Pteridophytes and Phanero-

gams the plant body is covered by a distinct outer layer of cells or
EPIDERMIS; this is wanting in plants of the lower groups. A
typically developed epidermis consists of one layer of cells (Fig. 74 e),
the outer walls of which on the free surfaces of the plant are more
strongly thickened. This is especially the case in all aerial parts of
plants adapted for a long life, but on the more perishable parts of a
plant, such as the floral leaves, or on subterranean parts such as the
roots, the cells of the epidermal layer are generally thin-walled or
only slightly thickened. For such reasons the epidermis of the root
was formerly termed the epiblem. Even when the external walls
of the epidermal cells are considerably thickened, the side walls, at
least in part, remain unthickened. The external walls are also more
or less cuticularised, while their outermost layer, which is more
decidedly cuticularised and capable of withstanding even the action
of concentrated sulphuric acid, extends as a CUTICLE continuously
over the surface of the epidermis. This cuticle appears to be always
101
wanting from the epidermis of roots ( ). The cuticle has its origin
in the primary walls of the younger epidermal cells, which, during
the increase in size of the plant, grow in area, and at the same
time are strengthened by the deposition of cutin. The cuticle of
leaves in the tropics is often specially smooth and shining. It reflects
a portion of the sunlight, giving rise to the sparkling appearance
characteristic of tropical foliage, and serves to protect the plant from
too strong insolation. The cuticle frequently becomes folded, and so
assumes a striated appearance (Fig. 114). By the thickening and the
cutinisation of their outer walls the mechanical rigidity of the epi-
dermal cellsis increased, and the loss of water by transpiration is

lessened. Plants in dry climates, or so situated that, for any reason,

transpiration from their outer surfaces must be diminished, are char-
acterised by the greatly thickened and cuticularised walls of their epi-
dermal cells. In some of the Gramineae, Equisetaceae, and many other
 MORPHOLOGY 101

plants, the cell walls of the epidermis are silicified. In the Equisetaceae
the impregnation with silica is so considerable that these plants are
used for polishing. Heating, even to redness, does not destroy the
structure of such silicified
epidermal cells.
Deposits of wax are also present in the cutinised layers of the
epidermis, and consequently water will flow off the epidermis
without wetting it. The wax is sometimes spread over the surface
of the cuticle as a wax This is the case in most fruits,
covering.
where, as is so noticeable on plums, it forms the so-called bloom.

Fio. 108. Transverse section of a node of the sugar-cane, Saccharum oflleinaru,m, showing wax-
incrustation in the form of small rods, (x 540.)

The wax coverings may consist of grains, small rods (Fig. 108), or
crusts.

The wax deposits attain their greatest thickness on the leaves of some of the
Palms on the Peruvian Wax Palm, Geroxylon andicola, the wax covering is more
;

than 5 mm. thick. This wax, as well as that obtained from the fruit of Myrica
cerifcra, is known as vegetable wax, and possesses an economic value. The wax
incrustations may be melted by heat they are soluble in ether and in hot alcohol.
;

In many cases, in place of the wax coverings, small grains and scales of a fatty
substance, which is soluble even in cold alcohol, are excreted. The dusty cover-
ings thus formed appear either mealy white or golden yellow, and are the cause of
the striking appearance of the Gold and Silver Ferns, especially in species of
Gymnogramme.
In many cases, slimy or sticky excretions are produced between
the thickening layers of the epidermis and the cuticle ; these press up
the latter and finally burst it. Such excreting surfaces often occur
on bud-scales. Sticky zones are frequently formed on stems, as in
the case of Lychnis viscaria and other Sileneae, as a means of protection
to the flowers higher on the stem from undesirable visitors. Small
creeping insects, which would otherwise rob the flowers of their honey,
102 BOTANY

seem as little able to pass beyond such a sticky zone, as other larger
animals to surmount the rings of tar often placed around the trunks
of fruit trees for a similar protective purpose. Excreting epidermal
surfaces form also the nectaries of flowers, which by means of their
sweet secretions attract such animals,
generally insects, as are instrumental
in their pollination.
The the epidermis are in
cells of

uninterrupted contact with each other,

and as a rule have undulating side
walls (Fig. 109). The protoplasm of
epidermal cells generally appears to
be reduced to a thin, peripheral layer,
and the sap cavities filled with a
Fid. 109. Surface view of the epidermis colourless or coloured sap. Around
from the upper side of a leaf of Mercuri-
their nuclei cluster the colourless rudi-
alis perennis. (x 300.)
ments of the chromatophores, showing
that, although exposed to the light, their further development into
chloroplasts may cease in cells not destined to take part in the
assimilatory processes. Such epidermal cells with undeveloped
chromatophores, besides acting as an external protection, serve as
water-reservoirs; their side walls, by means of folds in theunthickened
parts, can expand and collapse as a bellows, according to the varia-
tions in their supply of water. In plants which grow in shade
chlorophyll is usually pre-
sent in the epidermal cells.
The external walls of
the epidermal cells of this
upper surface of foliage
leaves not uncommonly
project or exhibit a lens-
shaped thickening in the
middle (Fig. 75) this ar-
;

rangement serves to con-

centrate the rays of light.
The refractivity of the
lens-shaped thickening is
FIG. 110. Epidermis with stomata from the lower surface
rendered greater by pec- of the leaf of Helleborvs niger. (x 120.)
tinisation, ctitinisation,
silicification, or waxy secretions. In the cases where the optical
apparatus is formed by a bulging of the uniformly thickened wall, the
1C2
cell-sap containing much tannin serves as the refractive medium ( ).
103
The formation of stomata ( )
in the epidermis is characteristic
more highly-developed plants which are exposed to
of all parts of the
the air. Each stoma consists of an intercellular passage perforating
the epidermis and bounded by two elliptical epidermal cells, termed
 MORPHOLOGY 103

GUARD-CELLS (Figs. 110, 111 A). The guard-cells always contain

chloroplasts, and are also characterised by their peculiarly thickened
walls. best seen in transverse sections, form ridge-like
These, as is

protuberances projecting above and below from the sides of the

Fid. 111. Epidermis from the under side of a leaf of Tradescantia virginiai. A, In surface view ;

B, in transverse section I, colourless rudiments of chromatophores surrounding the nucleus.

;

(X 240.)

guard-cells adjoining the air-passage (Fig. Ill B). Midway between

the projecting ridges, on the other hand, the walls of the guard-cells
remain unthickened (Fig. 112).

The unthickened parts of the walls of guard-cells jut out into the pore (Figs.
Ill 112), and thus facilitate its closing.
,
In addition, the external thickened

FIG. 112. Transverse section of the epidermis of Aloe nigricans. i, Inner, uncutinised thickening
layer, (x 240.)

walls of the two adjacent epidermal cells become, in some cases, suddenly thin on
approaching the guard-cells. By this means a hinge-like connection is formed
which renders the guard-cells more or less independent of the other epidermal cells.
At other times the same result is accomplished by raising the stomata above the
epidermis, or, more frequently, by sinking them below the less thickened epidermal
104 BOTANY PAHT i

walls. The elevation serves to increase the evaporation, and is on this account
met with in Ferns which grow in damp situations. The depression diminishes the
transpiration by creating a still atmosphere above the guard-cells. On this account
it ismet with in plants of dry regions (xerophytes). Frequently the epidermal
cellsadjoining the guard-cells are less thickened or lower than the other cells
of the epidermis (Fig. 111). Such special epidermal cells are called SUBSIDIARY
CELLS, and have the same use as the hinge joint mentioned above.
The stomata are formed by the division of a young epidermal cell into two cells
of unequal size, one of which, the smaller and more abundantly supplied with
protoplasm, becomes the stoma mother-cell while the larger, containing loss
;

protoplasm, usually forms an ordinary epidermal cell. The stoma mother-cell

becomes elliptical in outline and divides again, by a vertical wall, into the two
guard-cells, between which, by a splitting of the wall, the intercellular passage
(pore) is formed. Before the formation of the definitive stoma mother -cell, succes-
sive divisions of the young epidermal cell often occur ;
in such cases the finally

developed stoma is generally surrounded by subsidiary cells.

Stomata are chiefly developed on the green parts of plants, but

are sometimes found even on the coloured floral leaves. They are
naturally found in greatest numbers on the leaves, as it is there that
they are most needed to facilitate the interchange of gases in the
processes of assimilation. In dorsiventral leaves the stomata occur,
for the most part, if not exclusively, on the under surface, and

average about 100 to the square millimetre, although in some plants

their number may reach 700. Leaves which are alike on both sides
have their stomata equally distributed on the two surfaces. Floating
leaves of aquatic plants have stomata only on the side exposed to
the air. In some cases, as in the Oleander (Nerium Oleander),
numerous stomata are situated together in pit-like depressions of
the under surfaces of the leaves. In the tissue directly under each
stoma there is always a large intercellular space, termed the
RESPIRATORY CAVITY (Fig. Ill B), which is in direct communication
with the other intercellular spaces extending throughout the leaf
tissue. In plants grown in abundance of moisture, these intercellular
spaces are usually larger than in the case of plants growing in
drier situations.
In contrast to the stomata, which as air-pores serve for the
interchange of gases, a few plants also possess WATER-STOMATA or
WATER-PORES, situated at the ends of the veins or nerves of the
leaves. These pores serve as organs for the discharge of water or
watery solutions. Calcium carbonate, in solution, is frequently
excreted in this way, and in many species of Sdxifraga it forms white
scales on the margins of the leaves. Although water-pores may
often be found at the apices and tips of the marginal teeth of young
leaves, they seem to dry up as the leaves become more mature. The
guard-cells of water-stomata usually lose their living contents early,
and thus the passage between them remains continually open. The
water-stomata (Fig. 113) are always larger than the air-stomata.
 MORPHOLOGY 105

Although submerged leaves of aquatic plants are devoid of air-

stomata, water-stomata often occur on them.

Hairs or TRICHOMES and EMERGENCES are characteristic of the
tegumentary system. The cells of the hairs may retain their living
contents, or die and become
filled with air ;
in the latter
case the hairs appear white.
The simplest form of hairs are
the PAPILLAE, which are merely
epidermal cells, the external
walls of which have protruded
in a conical form. Papillae are
often developed on the petals
of flowers, and are the cause
of their appearance
velvety
(Fig. 1 14). hairs, such
Longer
as the root hairs (Fig. 170 r),
are also prolongations of single
epidermal cells ; these are
characteristic of definite
Fid. 118. Water-pore from the margin of a leaf of
regions of the root, and only Tropaeolum majus, with surrounding epidermal
a limited portion of the outer cells, (x 240.)

wall of the epidermal cell

protrudes as a hair. The woolly hairs found in young buds are
a protec-
generally similarly prolonged epidermal cells which, as
.

tive covering, surround the young growing tissues and some-

times remain on fully developed parts of plants to protect them
against too rapid evaporation
and direct insolation. The
hairs developed from some of
the epidermal cells of the seed
coats of various species of Gossy-
pium attain an unusual length,
and supply the cotton of com-
merce (Fig. 115). These cotton
FIG. 114. Surface of the upper epidermis of a petal hairs are Sometimes 6 Cm. long,
of Viola tricolor, showing ridge-like projections
f u |i y developed State
from the lateral walls, and protruding papillae.
an(j in
n n.
^^ . i

(x 250 )
contain only air ; their cell walls
are thicker than those of ordinary

hairs, and covered with a delicate

cuticle. They are usually some-
what flattened and at the same time twisted ; and are wider in the
middle than at either end. Only a certain number of the epidermal
cells of the seed grow out as cotton hairs (Fig. 115 BJ. BRISTLES
are short, pointed hairs, in the thickened cell walls of which calcium
carbonate or silica has been deposited (Fig. 116, below, to the right).
The STINGING HAIRS (Fig. 116), such as those of Nettles (Urtica)
106 BOTANY

and of the Loasaceae, are special forms of bristles, and arise as

prolongations of single epidermal cells. These, however, swell in

the course of their development, and becoming surrounded by
adjoining epidermal cells present the appearance of being set in
sockets ; while, at the same time, by the multiplication of the cells

FIG. 115. Seed-hairs of the cotton, Gossypium her- Fio. 116. Stinging hair of Urtica
baceum. A, Part of seed-coat with hairs (x 8) dioica,with a portion of the epi-
BI insertion and lower part, B% middle part, and dermis, ami, to the right, a small
83 upper part, of a hair, (x 300.) bristle, (x (X).)

in the tissue at their base, the whole hair becomes elevated on a

column-like protuberance. The hair tapers towards the apex and
terminates, somewhat obliquely, in a small head, just below which
the wall of the hair remains unthickened. As the wall of the hair is
silicified at the end and calcified for the rest of its length, the whole
hair is therefore- extremely stiff. Such hairs furnish a means of
defence against animals. The heads break off at the slightest touch,
and the hairs piercing the skin pour out their poisonous contents,
 MORPHOLOGY 107

which, especially in the case of some tropical nettles, may cause

severe inflammation.
The cells surrounding the base of a hair are often arranged in
a ring or in radiating lines, or are otherwise different from the
surrounding epidermal cells. Such cells may
be called subsidiary cells to the hairs.
UNICELLULAR HAIRS, such as we have
so far considered, may terminate in well-
defined heads resulting from the swelling of
their tips, or their side walls may develop
irregular excrescences ; on the other hand,
they may remain short and expanded like a
balloon, or remain close to the surface of
the epidermis as spindle-shaped or stellate
hairs. MuLTlCELLULAR HAIRS may be
merely simple rows of similar cells, as the
hairs on the stamens of Tradescantia (Fig.
60) ; or their terminal cells may become
FIG. 117. Glandular hair from
swollen into globular heads (Fig. 117), like
the petiole of Primula sinensis.
those on the Chinese Primrose (Primula (After DE BARY, x 142.)

sinensis) ; or an epidermis may be covered

with disc-, star-, or bowl-shaped hairs (Fig. 118). Sometimes the
hairs become variously branched, lose their living contents, and form
a silky or woolly protective covering similar to that formed by uni-
cellular hairs. In special cases, as in the
scale hairs of Ferns, they may even have
the shape of a small leaf.
EMERGENCES, unlike hairs, are not
formed solely by epidermal cells, but a
number of cells, lying more or less deeply
in the sub-epidermal tissues, also take part
in their formation. Thus, for example,
while only a few rows of sub-epidermal cells
enter into the formation of the emergences
FIG. 118,-Gianduiar scale from the (Fig. 119) on the margins of the stipules of
female inflorescence of the Hop, the Pansy (Viola tricolor}, much deeper- lying
Humvlus Lupulus, in vertical tisgue participates in the development of
section. A, before, B, after the
cuticle has become distended by the emergences which, as PRICKLES, serve
the excretion, in B the ex- j n t ne case o f Roses as a means of
protection,
cretion has been removed by
andj at the same tim e are
.
,1 , r ,

alcohol. (After DE BARY, x H2) of assistance in

climbing. The thick emergences, which

spring from the roots of the Podostemaceae, and serve to attach
them to rocks, are parenchymatous throughout, but vascular bundles
may be included within emergences, as is well shown in the club-
shaped digestive glands or tentacles (Fig. 120) on the leaves of the
Sundew (Drosera). Some emergences resemble in structure certain
108 BOTANY

of the metamorphosed members of the plant body described in the

preceding chapter ;
the resemblance between prickles and thorns, and
between haptera and lateral roots will serve
as examples. They are not, however, to be
traced back in origin to such members (p. 50).
Both hairs and emergences frequently
act as secreting organs, and are then termed
GLANDS. In many cases they are concerned
with the active exudation, and at times also
the absorption of water. They then belong
to the class of organs designated HYDA-
THODES ( 104 ) by HABERLANDT. Other glandular
hairs excrete 'a resinous substance. The
hairs of Primula sinensis (Fig. 117) are in

reality such glands, in

which the cuticle of the
terminal globular head
is pressed away from the

cell wall by the resinous

matter excreted from the
hair, until finally the
Glandular
Fir.. 110. colleter
bulging CUticle IS ruptured
from a stipule of Viola trl-
an(j the res inOUS Secretion
color, showing also a uni- . .

cellular hair, (x 240.) exudes. 1 he hairs of this

and other species of Pri-
mula (especially P. obconica) are capable of excit-
ing inflammation in the skin of those handling
them ( 10C ). Only some persons are susceptible
to the effect. The similar but more complicated
glandular hairs of Hops (Fig. 118) produce a
secretion called LUPULIN, to which beer owes its
bitter taste and distinctive aroma. The secretion
is set by the bursting of the cuticle, the
free
latter having been previously raised up from the
underlying cell wall as a continuous membrane
(Fig. 118 B}. Hairs and emergences with abund-
ant protoplasmic contents occur on irritable
stamens, perianth leaves, and pulvini, and, as
HABERLANDT 105
) showed, act as tactile papillae,
(

hairs, or bristles in the reception of stimuli. In

other cases they are not themselves irritable, but FIG. 120. Digestive gland
from Drosera rotu " fli -
serve to conduct a stimulus mechanically towards
folia, (x 60.)
the irritable tissue.

The mucilaginous matter produced

in young buds by the mucilage papillae or
COLLETERS results from the partial dissolution of the cell wall under the cuticle.
After the mucilaginous secretion has been discharged by the ultimate rupture of
 MORPHOLOGY 109

the cuticle, another new cuticle forms over the cell wall, and the process is again
repeated. The colleters are special forms of hairy structures, and are often de-
veloped in buds to protect the young organs from drying, by means of the
mucilaginous modification of their cell walls. Where the dissolution of the cell
wall is accompanied by secretions from the underlying cells, the colleters assume
rather the character of glandular hairs. Such GLANDULAR COLLETERS are common
in the winter buds of trees in the Horse-chestnut (Aesculus ffippocastanum), for
;

example, the bud-scales of the winter buds are stuck together by a mixture of
gum and resin, which has exuded from colleters of this nature. The glandular
hairs of the Pansy (Fig. 119) act in a similar manner. The emergences on the
leaves of the Sundew (Drosera), described as digestive glands (Fig. 120),
discharge
glistening drops of mucilaginous matter, not under the cuticle, but from the free
surface of the glands at the ends of the tentacles. Small animals are caught by
means of these sticky excretions, and are afterwards digested by the plant. The
nectaries also often excrete sugary solutions directly from their surfaces. In
flowers these serve to attract insects, which effect pollination, while on other parts
of the plants they are known in certain cases to attract ants, which protect
the plant. The osmotically active substances in the nectar are in the first
instance derived by transformation of the outer cell walls, or are secreted by the
cells. The presence of these substances on the surface of the nectary attracts
water from the tissue beneath, and thus leads to the continued formation of the
nectar.
In some of the Piperaceae and Begoniaceae, and in some species of Ficus, the
epidermis is composed of several
layers but this is a comparatively rare occurrence.
;

Such a many-layered epidermis results from a division of the young epidermal

cells parallel to their external surface. The epidermis of Ficus elastica (Fig. 76)
has three layers, and serves as a reservoir for accumulating water. The cystoliths
of Ficus elastica, already referred to (p. 63), occur in single swollen epidermal cells.
A many-layered root epidermis is also met with, as in species of Asparagus, Crinum
and Lycoris ( 107 ). The many - layered epidermis of the aerial roots of many
Orchids, and of various Aroids, undergoes a peculiar modification and forms the
so-called VELAMEN RADICUM (p. 48), a parchment-like sheath surrounding the
roots, and often attaining a considerable thickness. The cells of this enveloping
sheath are generally provided with spiral or reticulate thickenings, and lose their
living contents. They- then become filled with either water or air, depending upon
the amount of moisture contained in the surrounding atmosphere. These root-
envelopes absorb water like blotting-paper when the velamen is filled with water,
;

the underlying tissues impart a greenish tint to the root but if it contains only
;

air the root appears white. The epidermis of fruits, and particularly of seeds,
exhibits a considerable variety of modifications in its mode of thickening, and in
the relations the thickening layers bear to one another. The purpose of these
modifications in the epidermis becomes at once evident, when it is taken into
consideration that, in the case of fruits and seeds, in addition to protecting and
enclosing the internal parts, the epidermis has often to provide for their
dissemination and permanent lodgment.

The Vascular Bundle System. The PRIMARY VASCULAR BUNDLES

extend in the form of strands throughout the body of the higher
plants. In more transparent stems, such as those of Impatiens parvi-
flora, bundles may be clearly distinguished and their course
the
followed. The arrangement of the bundles of leaves is apparent from
110 BOTANY

the venation. In many parallel-veined leaves the bundles are easily

isolated. This is often done accidentally, as when, for example, in
picking a leaf of Plantain (Planfayo media) a pull is given at the same
time.
Special strands of tissue serving for the transport of substances
through the plant are found in the more highly differentiated Thallo-

Fio. 121. Transverse section of a vascular bundle from the internode of a stern of Zea Mais, a, King
of an annular tracheid ; sp, spiral tracheid n
and m', vessels with bordered pits v, sieve-
; ;

tubes ; s, companion cells cpr, compressed protophloem I, intercellular passage ; vg, sheath
; ; ;

/, cell of fundamental tissue, (x 180.)

phyta examples are afforded by some of the lied and Brown Seaweeds
;

(Rhodophyceae and Phaeophyceae). In the Laminariaceae these con-

108
ducting tracts contain elements which closely resemble sieve-tubes ( ).
The thallus of some Liverworts is traversed by a strand which
resembles the nerve of a leaf. Bundles sharply limited from the
surrounding tissues first appear in the Mosses they occur commonly ;

in the leaves, less often in the stems. A fairly simple example of

this kind of conducting bundle is that of the stem of Mnium
which is
represented in tranverse section in Fig. 161.
 MORPHOLOGY 111

Such bundles reach their highest differentiation in the Polytrichaceae.

In them the stem has a central cylinder composed of elongated
cells with scanty contents, of elements resembling sieve-tubes, and of

elongated cells ; the three kinds of element serve respectively to

109
transport water, albuminous substances, and carbohydrates ( ).
The arrangement of the tissue of these complicated strands is often
similar to that found in the vascular bundles of more highly organised
plants. This is a good example of a striking resemblance between

*p f /*
I

FIG. 1-2-2. Longitudinal section of a vascular bundle from the stem of Zea Mais, a, and a', Rings of
an annular tracheid v, sieve-tubes; s, companion cells; cp, protophloem I, intercellular
; ;

passage vg, sheath sp, spiral tracheides. (x 180.)

; ;

structures which are analogous but have arisen independently in the

course of evolution.
It is, however, in the Cormophytes, which possess roots, that a

high degree of differentiation of the vascular bundles is first attained.

Since the absorption of water is limited to the roots, the arrange-
ments in the conducting tracts require to be more perfect. Two
distinct components can be distinguished in these vascular bundles,
the TRACHEAL Or XYLEM PORTION, and the SIEVE-TUBE or PHLOEM
PORTION. While these may form independent strands, they are gener-
ally united inone VASCULAR BUNDLE. Other terms often used to
designate the vascular bundles are FIBRO-VASCULAR BUNDLES and
MESTOME. The vascular portion is also termed the XYLEM or
112 BOTANY FART 1

no
HADROME, and the sieve-tube portion the PHLOEM or LEPTOME ( ).
The distinction of the two components of the vascular bundle is
most evident from transverse sections (Figs. 121, 123), with which
the longitudinal section (Fig. 122) should be compared. The
vascular portion contains TRACHE/E and TRACHEIDES as most essential
for the fulfilment of its function of water conduction (a, sp, m, Figs.

Fio. 123. Transverse section of a vascular bundle from a stolon of Ranunculus repens. s, Spiral
tracheides ; m,
vessel with bordered pits c, cambium ; v, sieve-tubes ; vg, sheath, (x 180.)
;

121, 122), or tracheides alone, and, in addition, living, elongated

parenchymatous cells that may be designated XYLEM or WOOD
PARENCHYMA. In the phloem portion the most essential elements
(#), which serve for the conveyance
are the SIEVE-TUBES of albuminous
matter. They are always accompanied by other living cells ; either
by the so-called COMPANION CELLS (s), or in addition by elongated
parenchymatous cells, or by the parenchyma alone. Companion
cells only occur in the phloem of Angiosperms. They are sister
 MORPHOLOGY 113

cells of the sieve-tubes, having arisen by longitudinal division

from the same mother cell. The companion cells are not so large as
the sieve-tubes, and may be distinguished from them by their more
abundant protoplasmic contents, and especially by the fact that they
retain their nuclei, while the nuclei of the sieve-tubes soon disappear.
In Monocotyledons (Figs. 121, 122), and in the Ranunculaceae among
the Dicotyledons (Fig. 123), the phloem consists solely of sieve-tubes
and companion cells ; in the other Dicotyledons parenchymatous
elements are also present, and these are accordingly distinguished
as PHLOEM PARENCHYMA no companion cells are found in Gymno-
;

sperms and Pteridophytes, and

-

in addition to sieve-tubes the phloem

contains only phloem parenchyma.
The bundles of the Phanerogams (Gymnosperms and Angiosperms)
are generally COLLATERAL in structure, that is, the xylem and phloem
are in contact on one side only. In stems the most usual arrange-
ment of the two portions of a collateral bundle is that in which the
xylem lies nearest the centre in leaves the xylem portion lies nearer
;

the upper, and the phloem portion nearer the lower surface. Closely
allied to the collateral type is the bicollateral type of bundle. In
this the xylem is accompanied by phloem both on the outside and
inside. Such bicollateral bundles are characteristic of the Cucur-
bitaceae ( 1U ). The xylem and phloem of roots generally form
separate strands (Fig. 124 s, v), and the xylem strands are differently
oriented ; while in stems the narrow vessels are nearer the centre
and the wider nearer the circumference, in roots this order is exactly
reversed.
The "CONCENTRIC" vascular bundles of the Pteridophyta (Fig. 125)
contain tracheides (sp), and only in exceptional cases tracheae (sc).
The latter are as a rule wanting in Pteridophyta, although this
group goes by the name of Vascular Cryptogams. The water-con-
ducting elements exhibit scalariform thickenings, only the narrowest
having spiral markings (sp) ; they are surrounded by xylem paren-
chyma (Ip). Outside this comes a zone of tissue consisting of
sieve-tubes (v) and phloem parenchyma (s).

A number of similar vascular bundles are present in the stem of most Ferns and
species of Sclaginella. In Lycopodium they are fused into a single central cylinder.
In the stem of Equisetaceae vascular bundles of collateral structure appear.

The vascular bundles are developed from strands of meristematic

tissue which are called PROCAMBIUM STRANDS. Within each strand
a zone of tissue commences to divide tangentially, and behaving
as a primary meristem produces to both the inside and the outside
new cells in radial order. If the whole meristematic tissue of a
procambium strand is exhausted in this process, the vascular bundles
are said to be CLOSED but if any of the meristematic tissue remains
;

in an undifferentiated condition between the xylem and phloem

I
114 BOTANY I'AKT 1

portions, the bundles are spoken of as OPEN (Fig. 123 c). The
Pteridophytes have, almost without exception, closed bundles ;
in

Monocotyledonsalso the bundles

are always closed (Fig. 121);
Gymnospermsand Dicotyledons
(Fig. 123), on the contrary, have
open bundles.
In those portions of plants which
are still actively growing in length,
the procambium strands remain un-
differentiated, except at definite

points, where single rows of cells lose

their meristematic condition and form
narrow, annular, and spiral vessels
and sieve-tubes, or sieve-tubes and

companion cells the structure of all of

;

these is of such a nature as to render

their elongation possible. Such prim-
ary vascular elements are termed
while the correspond-
Fio. lL'4. Transverse section of central portion of . . , ...

the root of Aeenu Calamus, m, M'-dnlla ; .--,

6 sieve elements are in like manner
ing
xylem; r, phloem ; p, pericycle ; c, endodermis ; designated PROTOFHLOKM. The pro-
c, cortex, (x 90.) .
toxylem occuj>ies the innermost, the
protophlocm the outermost side of a
procambium strand, from which a collateral bundle is eventually formed. After
the growth in length of any part of a plant ceases, the differentiation of the pro-
cambium strand into a collateral vascular bundle is continued from the inner and
outer sides of the strand toward the centre.
In fully developed vascular bundles the protoxylem and protophloem cease to
perform their functions. The protoxylem elements become compressed and
ruptured by the tension resulting from the continued vertical growth (a and a',
Fig. 122), so that in their stead a lysigenic intercellular space is often formed
(Figs. 121, 122). The protophloem elements (cf. Figs. 121, 122) at the same time
become disorganised, and their sieve-plates closed by a covering of callus.
In accordance with the inverted orientation of the xylem, the protoxylem of
roots is found on the outer, not on the inner side of the vascular strands (Fig. 124).

The Terminations
of the Vascular Bundles. In leaves,
particularly in the foliage leaves of Angiosperms, the vascular bundles
become much branched until finally they are reduced to extremely
fine strands. In the leaves of Gymnosperms this branching of the
bundles does not usually take place, but instead, a single vascular
bundle frequently runs throughout the whole length of the leaf.
The vascular bundles of the reticulately-veined leaves of Dicotyledons
illustrate the most extreme form of branching.

The minute distribution of the bundles in the leaf-lamina facilitates the regular
conduction of water to all parts of the leaf-tissue, and at the same time renders
easier the removal of the assimilated products. In the same degree as the ramifica-
 MORPHOLOGY 115

tions of the vascular bundles are continued, the bundles themselves become
attenuated and simpler in structure (Fig. 126). The vessels first disappear, and
only spirally and reticulately thickened tracheides remain to provide for the water
conduction. The phloem elements undergo a similar reduction. In Angiosperms,
in which the sieve-tubes are accompanied by
companion cells, the sieve-tubes become
narrower, whilst the companion cells retain their original dimensions. Finally,

FIG. 125. Transverse section of a concentric bundle from the petiole of 1'teris aquilina. 'sc,
Scalariform vessels ; sp, protoxylem (spiral tracheides) ; sc*, part of a transverse wall showing
.scalariform perforations ; Ip, xylem parenchyma v, sieve-tubes
; ; j>r, protophloem ; pp, starch
layer; e, endodermis ; s, phloem parenchyma, (x 240.)

in the cells forming the continuation of the sieve-tubes, the longitudinal division
into sieve-tubes and companion cells does not take place, and TRANSITION CELLS are
formed ( 112 ). With these the phloem terminates, although the vascular portion of
the bundles still continues to be represented by short spiral tracheides. The
ultimate branches of the bundles either terminate blindly or anastomose with other
vascular bundles.

The Fundamental Tissue System usually forms the main bulk

of the primary tissues of the body of a plant. The whole tissue of
the lower plants, as it shows no internal differentiation, may, in a
certain sense, be considered fundamental tissue. The other tissues
116 I JOT ANY

have gradually arisen from the fundamental tissue in the course

of phylogenetic development. The fundamental tissue in the
higher plants is enclosed by tegumentary tissue, and traversed 1>v
the vascular bundle system. While the tegumentary tissue protects
the plant externally, and the vascular bundle system performs the
office of conduction, and also of mechanically strengthening the

plant, the duty of providing for the nutrition of the plant and of
storing reserve food material falls chiefly to the fundamental tissue.
The fundamental tissue consists, therefore, for the most part of
parenchymatous containing chloro-
cells

phyll, at least to such depth as the

light penetrates; internally a colourless
parenchyma is found. The fundamental
tissue system also takes part in pro-
viding for the mechanical rigidity of
plants, and in connection with this
function it possesses collenchyma (Fig.
74 c) and sclerenchyma as its special
mechanical tissues. -The COLLENCHYMA
(p.68) is unlignified and very elastic, and
thus fitted for stretching ; it is the form
of mechanical tissue suitable for those
parts of plants still undergoing growth
in length. The sclerenchymatous fibres,
which are often unlignified but have
greatly thickened walls, on the other
hand, are formed after growth in length
has ceased, and sclereides (p. 73) arise
FIG. 126. Termination of a vascular even later. The elongated cells of the
bundle in a ,leaf of Imputiens po,rvi
fundamental tissue also perform a certain
flora, (x 240.)
share of the work of conduction, and
serve for the transport of carbohydrates. Secondary or waste pro-
ducts, resulting from chemical changes, are also deposited in special
cells of the fundamental tissue. Consequently idioblasts (p. 73),
containing crystals or rows of crystal-containing cells, are often met
with in the fundamental tissues, together with cells, tubes, cavities, or
canals containing tannin, gum, resin, ethereal oils, latex, or alkaloids.
Such waste products are for the most part deposited near the surface
of a plant, in order to serve as a defence against destructive animals,
or that they may afterwards be thrown off along with the superficial
tissue. Cells containing these waste products, particularly crystal
cells and latex tubes, are often found, accompanying the phloem

portion of the vascular bundles.

The Aroideae, Nymphaeaceae, and several other plant families possess a
peculiar form of idioblast, in the so-called internal hairs, which project into the
intercellular spaces of the fundamental tissue. In the wide intercellular passage*
 MORPHOLOGY 117

'ofthe petioles and flower stems of the Water-Lily these idioblasts are stellate in
form. Their walls are strongly thickened, and provided with short protuberances
in which small crystals of calcium oxlate are
deposited.

The Distribution of the Primary Tissues (

m )

In the body of multicellular plants a distinction between an outer

small-celled and firm tissue and an inner large-celled looser tissue
soon becomes apparent. The outer tissues are best adapted for pro-
tection, the inner for conduction and storage. The cells of the inner
tissues accordingly become elongated for the purpose of conduction.
The outer tissues in plants, which must provide independently for
their own nourishment, contain
chromatophores fitted for assimi-
lation, and are correspondingly
coloured, while the inner tissues
remain colourless. The outer
portion of the fundamental tissue
thus differentiated is called the
CORTEX, the inner the MEDULLA
or PITH. An epidermis, distin-
guishable from the cortex, is found
in some of the Mosses, but a sharp
distinction between these tissues
is first found in the more
highly
organised plants.
In the Stem of a Phanerogamic
plant there is an outer skin or
epidermis (Fig. 128 B, e) on the FK-. I-JT. Tr:msv.-i-s>- sirti.m ui .m mtMiio<ie of
th " sti * p r ilniiry c rte x :
external surface ; then follows the
'

pc,
'f :"'
^ ?
pencycle co vascular bundles go, fuuda-
;
,
, ,
;

PRIMARY CORTEX (Figs. 127, 128 mental time of the eentnl eyliwter. (x 2.)
A, pr), and internal to this the
so-called CENTRAL CYLINDER, for which VAN TIEGHEM has proposed
the name STELE (column) ( 114 ). The innermost layer of the primary
cortex, which may be designated by the term PHLOEOTERMA, is for
the most part not distinctly differentiated, but can be recognised in
the aerial stems of land-plants as a starch-sheath while in the ;

rhizomes of land-plants and in the stems of water-plants it forms the

ENDODERMIS. Differentiated as a starch-sheath (Fig. 128 A, B,
xt),
the phloeoterma is rendered conspicuous by the quantity of
movable starch contained in its cells. A starch -sheath is often
present in the young shoots, while it disappears or becomes limited
to certain parts of the older shoots (
llr<
).
When developed as an
endodermis, portions of the lateral walls of its cells become suberised
uo In a cross-section these suberised portions of the cell walls of
( ).
118 BOTANY I'AUT 1

the endodermis appear as dark spots (Fig. 130), but in a tangential

section as a wavy band. The CENTRAL CYLINDER of the stem contains
vascular bundles (cv), which, in the Equisetaceae, the Gymnosperms
and Dicotyledons (Fig. 128 A\ are arranged in a circle, whereas in
Monocotyledons (Fig. 127) they are irregularly distributed. In all
these cases the xylem portion of the vascular bundle is directed
towards the centre, and the phloem portion away from the centre of
the stem. That part of the peripheral tissue of the central cylinder
lying outside of the bundles is called the PERICYCLE (pc). If the

FIG. 128. A, Part of a transverse section of a young stein of Arittnliu-hiu xifiho. e, Epidermis jv, ;

primary cortex ; st, starch-sheath ; c, central cylinder pc, pericycle, in this case with a ring'of
;

sclerenchyma fibres ctf, phloem, and cv", xylem portions of the vascular bundle cb, cambium
; ;

ring; m, medulla; ins, primary medullary ray. (x 48.) B, Small portion of the periphery
of a similar section of a still younger stem, e, Epidermis pr, primary cortex st, starch-
; ;

sheath with easily movable starch grains pc, outer layers of the pericycle. (x 350.)
;

bundles are arranged in a circle (Fig. 128 A\ that part of the

central cylinder enclosed by them is the PITH or MEDULLA (m), and
the tissue between the different bundles the PRIMARY MEDULLARY
RAYS. In the case of scattered bundles (Fig. 127), a distinction
between medulla and medullary rays is no longer possible. Wherever
there is no sharp distinction between primary cortex and central
cylinder, comparative investigation alone can determine whether a
tissue belongs to the primary cortex or to the central cylinder.

Although the fundamental tissue of the primary cortex is mainly a

chlorophyll-containing tissue, portions bordering on the epidermis frequently become
converted, for mechanical purposes, into strands of collenchyma or sclerenchyma.
Such a mechanical tissue, which serves to strengthen the epidermis, is known as a
HYPODKKMA. Of the tissues composing the central cylinder, the pericycle, the
 MORPHOLOGY 119

primary medullary rays, and medulla consist of fundamental tissue, and are chiefly

composed of colourless paren-

chyma. A part, however, of
the tissue of the pericycle
may become sclerenchymatous
(Fig. 128 A, pc) ; sclerenchy-
matous elements also often
surround individual bundles as
sheaths, or accompany the
phloem portion in the form of
strands (Figs. 121, 123). When-
ever such a sheath of scleren-
chyma developed about a
is

bundle, is
it interrupted on
both sides of the bundle, at
the junction of the xylern and
phloem portions, by paren-
chymatous cells, or by cells
which are only slightly thick- FIG. 129. Transverse section of an adventitious root of

ened and lignitied. The exist-

Allium Cepa. ep, Remains of the epidermis ex, exo-
;

dermis c, primary cortex

; e, endodermis
; <x, central
;
ence of these unthickeiied places
cylinder, (x 45.)
facilitates the exchange of water
and food material between the vascular bundles and the fundamental tissue. When

FIG. 130. Transverse section of an adventitious root of AlHitm Cepn. r, Primary cortex c, endo- ;

dermis ; p, pericycle; a, annular tracheides sji, spiral tracheides; sc and sc* ;

; scalarifonn
vessels; v, phloem, (x 240.)

a common starch-sheath is not present in a stem, starch-sheaths are sometimes

120 BOTANY

found surrounding the individual vascular bundles, or particular rows of cells

provided with movable starch grains are present. The central cylinder of
Phanerogams is simple, and occupies a more or less central position. In a few
cases, as in the stems of Auricula and Gunnera, the central cylinder is broken
vip into several partial cylinders.

In Roots, the division between primary cortex and central cylinder

is sharply marked by the endodermis, into which the innermost layer
of the primary cortex is usually transformed (Figs. 124, 129, 130 e).
The central cylinder becomes completely shut oft' from the primary
cortex by the suberisation of the
lateral walls of the endodermal cells,
and by their close and uninterrupted
contact. While, by this means, the
passage of gases from the intercellular
spaces of the cortex into the central
cylinder, with the consequent obstruc-
tion of the water -channels, is pre-
vented, the passage of water from the
cortex to the central cylinder can, at the
same time, go on unhindered through
the unsuberised inner and outer walls
of the endodermal cells. In this
manner it is possible for the water,
absorbed from the soil by the root-
hairs or by the surface of the roots, to
Fir,. i3i.-i>art of a transverse section of \>Q transferred to the tissues of the
a root of [rin Jlorentina. e, Endodermis, , i-
central cylinder.
j T ^v. u
In the older parts
showing ceil walls thickened on one
side.; /, transfusion cell; p, pericycle ; of the TOOtS, which no longer absorb
/; phloem vessel of xylem cortex.
; s,
water from the soil, the cells of the
; c,

(X 240.)
endodermis become greatly thickened,
but generally on one side only ; they may also be cutinised.
Should thickening occur at an early stage, special endodermal
cells, directly external to the xylem strands, remain unthickened
and serve as TRANSFUSION CELLS (Fig. 131 /).
While the root-hairs are as a rule developed from the cells of a definite region
of the epidermis, they may, in case the epidermis is thrown off at an early >t;igc,
arise from the outermost cortical layer, which then assumes the functions of an

epidermis. In any case the epidermis soon disappears, and the outermost cortical
layer becomes cuticularised and, as an EXODKKMIS, takes its place. Frequently
some of the cells of such an exodermis remain uucutinised and serve as transfusion
cells. They may be characterised their smaller size, and be regularly
by
distributed between the cutinised cells.In aerial roots the epidermis usually
forms a many-layered root-sheath or velamen (cf. p. 109), within which comes
the exodermis.

The primary cortex of the root is composed of colourless tissue,

which is
usually parenchymatous. In the outer layers the cells are
 MORPHOLOGY 121

in close contact with one another, but intercellular spaces are present
more internally. These intercellular spaces often widen into air-
cavities or passages. In many roots ahypoderma giving mechanical
support to the epidermis or exodermis is present. The outermost
layer of cells of the central cylinder (Figs. 124, 131 p) forms the
pericycle, which is also called the pericarnbium ; this is usually a single
layer, and in rare cases is wanting. The xylem and phloem portions
form separate strands (p. 113), radially disposed and alternating
with each other (Figs. 124, 130). It has already been shown that
the narrowest elements of the vascular strand are outermost. Roots
are described as diarch, triarch, polyarch, according to the number
of the vascular strands. For example, the roots of Acorns
Calamus (Fig. 124) are octarch, those of Allium Cepa (Fig. 130)
hexarch. The vascular strands may eitHer meet in the centre
(Fig. 130), or they may surround a central pith (Fig. 124).
The Leaves are composed of fundamental tissue (which is here
termed mesophyll), bounded by an epidermis and traversed by
vascular bundles. Sheaths are present around the bundles, extend-
ing to their fine terminations. The cells composing these mesophyll
sheaths are as a rule elongated and not separated by intercellular
spaces. Besides limiting the vascular bundles from the mesophyll,
the sheaths perform the important function of conducting soluble
carbohydrates from the leaf to the stem. The larger vascular
bundles are usually accompanied by strands of sclerenchyma ; these
disappear from the finer branches. Other strands of sclerenchyma
not connected with the vascular bundles may also occur in the
mesophyll and contribute to the rigidity of the leaf.

The mesophyll passes into the primary cortex of the stem, while the vascular
bundles are continuous with the central cylinder. Thus in the leaf, tissues corre-
sponding to the cortex and central cylinder of the stem remain distinct from one
another.

The mesophyll of the coloured FLORAL LEAVES of the Angiospei'ms

usually consists of a somewhat loose tissue, containing intercellular
spaces and traversed by vascular bundles. The laminae of many
assimilating FOLIAGE LEAVES, especially of shade-loving plants, may
have a similar uniform structure ; but they are usually more com-
plicated, and exhibit a difference in the structure of their upper and
lower sides (Fig. 132). In such dorsiventral structures the upper
epidermis is succeeded by one or more layers of cylindrical
parenchymatous elements elongated at right angles to the surface, and
known as the PALISADE CELLS. These are especially rich in chlorophyll,
and are often separated laterally from one another, so that the con-
duction of substances is limited to the direction of their longer
axis (Fig. 132). Adjoining the palisade parenchyma, and extending
to the epidermis (ep") on the under surface of the leaf, is a loose
122

tissii SPONGY PARENCHYMA. In contrast to the palisade

called the
cells, the the spongy parenchyma are less abundantly
cells of

supplied with chlorophyll they are also much more irregular in

;

shape, and have large intercellular air-spaces between them. The

palisade cells are elongated in the direction in which the rays of
light penetrate the leaf-lamina, and by this means are particularly
adapted to their special function of assimilation. The spongy paren-
chyma, on the other hand, is arranged to facilitate the free passage of
gases, and to that end develops large intercellular spaces in direct
communication with the stomata of the lower epidermis. HABER-
LANDT (
m )
has estimated that to every square millimetre of surface
in a leaf of Ricinus communis there are, in the palisade cells, 403,200

FIG. 132. Transverse section of a leaf of Vagus sylvaticc. ep, Epidermis of upper surface ; ep",
epidermis of under surface ; ep"', elongated epidermal cell above a vascular bundle ; pi, palisade
parenchyma ; s, collecting cells sp, spongy parenchyma ; k, idioblasts with crystals, in k'
;

with crystal aggregate; st, stoma. (x 300.)

chlorophyll granules ; in the cells of the spongy parenchyma only

92,000 ; that is, 82 per cent of all the chlorophyll granules belong to
the upper surface of the leaf, and only 18 per cent to the under side.
The palisade cells are often arranged in groups, in which the lower
ends of the cells of each group converge (Fig. 132). In this way
several palisade cells come into direct contact with a single expanded
cell of the spongy parenchyma, which thus functions, apparently, as
a collecting cell for a group of palisade cells. The products of
assimilation are passed on from the collecting cell through the
spongy parenchyma, to be finally carried to the mesophyll sheath
surrounding the vascular bundles. The sheaths serve as a
conducting tissue towards the stem.
At the base of the lamina the tissues close together and pass into
the leaf-stalk, where one is present. The dorsiventral structure
becomes less marked in the petiole. The cells are mostly elongated
SECT, i MORPHOLOGY 123

in the length of the petiole, a, modification which facilitates the con-

duction of food material. They are often thickened and so arranged
as to meet the altered requirements for mechanical rigidity.
The vascular bundles pass from the leaf-stalk into the stem and
there either arrange themselves among the bundles of the central
cylinder or at once fuse with some of them. In the leaf-stalks of
Angiosperms the bundles usually appear arranged in a curve open
above, but may form other figures. In the petioles of Ferns, the
partialcylinders are accompanied, as in the stem, by sclerenchy-
matous fibres forming strands or plates. It is the peculiar arrange-
ment of those brown-walled sclerenchymatous masses which forms
the double eagle apparent on cross-sections of the petiole of Pteris
from which the plant derives its specific name.
aquilina,

In certain families of the Dicotyledons, particularly iu the Crassulaceae, the

mesophyll of the leaf-lamina forms peculiar masses of tissue called the EPITHEMA
between the swollen terminations of the bundles and the epidermis. The cells of
the epithema are small and, for the most part, devoid of chlorophyll ; they are full
of water, and joined closely together, leaving only very small interspaces, which are
filled with water. They are internal hydathodes (cf. p. 108) and serve for the active
excretion of water. Water pores are usually situated above such epithemata. The
leaf-tips of a number of aquatic Monocotyledons show a depression into which the
terminations of tracheides project. These depressions arise by the destruction of
water-pores or of these together with the epidermis. They may be closed by the
persisting cuticle. These apical openings serve in the same way for the excretion
of water ( 118 ).

The Course of the Vascular Bundles ( 119 ). The bundles exhibit

a definite course and arrangement within the body of a plant. It is
sometimes possible, by maceration, to obtain preparations in which the
course taken by the bundles may be followed. By allowing a leaf,
stem, or flower to lie in water until it has become softened and
disintegrated, a skeleton formed by the more imperishable vascular
system may be obtained.
Vascular bundles which pass from a leaf into a stem form within
the latter what are known as LEAF-TRACES. The leaf-traces may be
composed of one or more vascular bundles, and are accordingly dis-

tinguished as one-strand or many-strand leaf -traces. When, as is

usually the case, the vascular system of the stem is entirely composed
of leaf-traces, each vascular bundle of the trace after passing down-
wards for some distance unites with another entering from a lower
leaf. The arrangement of the bundles in a stem varies according to
the distance and direction traversed before the coalescence of the
bundles takes place. A relatively simple case is afforded by the
young twigs of the Dwarf Juniper (Juniperus nana) (Fig. 133). The
leaves are in whorls of three, the leaves of successive whorls alter-
nating with one another. From each leaf a leaf-trace consisting of a
single vascular bundle enters the stem. This divides into two about
124 BOTANY

the middle of tbe internode below, and the divisions diverge and
unite with the leaf-traces of the whorl below. The arrangement of
the bundles may be shown diagrammatically by representing the
bundles as if on the surface of an unrolled cylinder, so that they all
appear in one plane. This is done in Fig. 133, which also shows the
origin of the vascular bundles of the axillary shoots (k).
The arrangement of the bundles iu the Yew (Taxus laccata),
although its leaf-traces have only one bundle, is much more compli-
cated (Fig. 134), for the bundles maintain a distinct course throughout
twelve internode 5 before coalescing. Each bundle at first descends
in a straight direction through four internodes ; it then curves to the
side to give place to a newly-entering
leaf-trace, with which it finally coalesces
at the twelfth internode. position The
of a leaf necessarily determines the
point of entrance of its leaf-trace into
the stem, and accordingly a diagram
(Fig. 134) of the bundles of Taxus will
exhibit a divergence of the leaf-traces
corresponding to the yV divergence of
the leaves. The course taken by the
in the stem, however, is
leaf -traces
independent
r of the leaf position, and
FIG. 133. Diagram of the course of tin-
vascular bundles in a young branch of
. .

varies considerably
, ,
,

m
.
,.
different stems,
jnniperuf nnnn shown on the unrolled although the divergence of their leaves
surface of the cylinder At I* the
vascular bundles passing to the axillary
be the game The gtem Q f Ckmaiis
/
shoots are seen. (After GEYI.KR.) vtiicella afford s an example of leaf traces

consisting of three vascular bundles.

The leaves are in whorls of two, the successive whorls alternating
with one another. The median strand of each leaf-trace (Fig. 135
ad, uk, nq, te) has a free course through one internode, and at the
node below divides into two arms which coalesce with the adjacent
lateral strands of the leaves inserted at this node. The two lateral
strands of each leaf-trace (Fig. 135 be, ef, hi, Im, op, rs) also are free
throughout the internode, but at the node below curve inwards and
become attached to the same lateral strands as the arms of the
median bundle of the trace.
The traces of the axillary buds of most Gymnosperms and
Dicotyledons unite to form two strands which enter the mother shoot
and are inserted upon its leaf-traces (Fig. 133 k).
As a general rule, the leaf-trace bundles in Gymnosperms and
Dicotyledons arrange themselves in a circle in the stem. There are,
however, Dicotyledons in which the vascular bundles form two
(Cucurbita, Phytolacca, Piper) or more circles (Amarantus, Papaver,
Thalidrum). In such cases the inner circle is usually more or less
irregular.
SECT. I MORPHOLOGY 125

In the stems of Monocotyledons (Fig. 127) the vascular bundles

are scattered, and without any apparent regular order. Their
scattered arrangement is due to the varying distances to which the
bundles of the leaf-traces penetrate into the central cylinder of the
stem. This results from the prolonged growth in thickness of the
12
growing point after the procambial strands have been laid down ( ).
A common arrangement of the bundles in monocotyledonous stems is
that of the so-called Palm type, in which each leaf-trace consists of

FIG. 134. Diagram showing the course of the vascular bundles in a shoot of Taius Imccvtv.

the numerous bundles which pass singly into the stem from the broad
leaf-base. The median bundle penetrates to the middle of the stem.
The depth to which the lateral bundles penetrate varies with their
remoteness from the median bundle. In their descending course the
bundles gradually curve outwards, and finally join other bundles
near the periphery of the stem. The number of internodes, therefore,
through which a bundle passes before coalescence, is variable the ;

median bundle, however, continues distinct for the longest distance.

The deeper penetration and greater length of the median bundle
become apparent in a median longitudinal section of such a stem
126 BOTANY PART 1

(Fig. 136). The numerous bundles entering the stem from axillary
shoots pursue a similarly curved course to those entering from leaves.
In addition to the leaf-
trace bundles or COMMON
BUNDLES, which are common
to both leaf and stem, there
are others, called CAULINE
BUNDLES, which belong solely
to the stem, and again others,
FOLIAR BUNDLES, which, Oil
entering the stern from the
leaf, at once coalesce with
other bundles and have no
independent existence in the
stem. The bundles of the
Pteridophytes are continued
as cauline bundles in the stem,

3/tl

nn

Fio. 135. Cieiiuitis viticella. End of a branch which

has been made transparent by the removal of the
superficial tissues and treatment with caustic
potash. The emerging strands have been slightly
displaced by gentle pressure. The two uppermost
pairs of young leaves (bli, W 2) are still without
FIG. 136. Diagram showing the
leaf-traces, r, Apical cone. (After NACF.LI.)
course of the vascular bundles
of Monocotyledons of the Palm
and those from the leaves join on to the
type, with alti-rriatiiix, two-
bundles of the stem. ranked amplexicaul leaves. The
numbers indicate the sequence
The stems of many Dicotyledons (Begonia, of the leaves m, median bundle.
;

Aralia) possess cauline bundles in addition to leaf- (After DE BARY.)

traces. The cauline bundles are situated in the
pith within the ring of leaf-trace bundles in the internode, and at the nodes are
connected by branches with the leaf-trace bundles.
 MORPHOLOGY 127

In the central cylinder of roots, the radially arranged strands of

xylem and phloem pursue a straight course parallel to one another.
The xylem and phloem strands of lateral roots are inserted upon the
corresponding strands of the parent root. Those of adventitious
roots are attached to the corresponding tissues of the stem or root
from which the adventitious root arises. The strands of xylem and
phloem of the main root of a phanerogamic seedling are continuous
with the leaf-trace bundles in the hypocotyl ( 12 ).
Increase in Thickness by Continued Enlargement of the Primary
Meristem. The growing points of those Monocotyledons which
have large stems (Palms, Pandanaceae, some Liliiflorae) exhibit this
feature. The growing point thus attains a considerable thickness, and
the stem has from an early period its definite circumference. The
increase in number of cells takes place in a zone lying at the
periphery of the growing point ( ).
m
A Special Form of Growth in Thickness of the Stem by means
of the Continued Enlargement of the Fundamental Tissue. This is
often exhibited by Palms, and leads in them to the formation of stems
which may be more than 1 metre in diameter. EICHLER ( 122 ) has
shown that the growth in thickness is solely due to the continued
expansion of the already existing cells of the fundamental' tissue of
the central cylinder. In this process, by the expansion of the cell
lumen and increased thickening of the walls, the strands of sclerenchy-
matous fibres accompanying the vascular bundles on their phloem
sides also become greatly enlarged. In this form of growth in
thickness no new elements are formed.
Enlargement due to Continued Cell Divisions in the Fundamental
Tissue. This is seen most strikingly in the growth of the fruits of
many Angiosperms. In their development from the small rudiment
in the flower there may be a great increase in size, although no new
meristematic tissue has been formed. The growth is due to repeated
divisions in the epidermal cells and in those of the fundamental tissue.
The tissue thus produced may when mature be thin-walled, or its cells
may undergo various changes in form and in the thickness of their
walls. From small structures such large fruits as those of the Gourds
rnay develop in this way.

123
The Secondary Tissues ( )

After growth in length is finished through the activity of a

cambial tissue, functioning either as a primary or secondary meristem
(p. 99), secondary tissues are added to the previously existing
primary tissues, or even substituted for them. Although, phylo-
genetically considered, secondary tissues seem to have been developed
first in the Pteridophytes in forms now only known in a fossil
128 BOTANY

condition (Caltimarieae, Sigillarieae, Lepidodendreae) they first became

of general occurrence in the Phanerogams. In them the formation
of secondary tissues is almost exclusively confined to the roots und
stems ; secondary growth is met with in some foliar structures but
only in a slight form.
124
Cambium ( ).
The primary and secondary meristems, the
activity of which gives rise to secondary growth, are given the name
of CAMBIUM. These tissues persist in a meristematic condition and
undergo successive divisions by walls parallel to one another, so that the
cells produced from them are arranged in radial rows. A similar mode
of division characterises the prim-

ary meristem in the procambial

strands, which give rise to col-
lateral vascular bundles (cf. p.
113), and this might also come
under the term cambium. It is

advisable, however, to restrict

this term to meristems giving
rise to secondary growth. As a
rule the cambium forms new
tissue-elements on both sides, but
cases are not wanting in which
this takes place to one side only.

As a
rule a single persisting initial
Fio. 137. Transverse section of a stein of Aristc
lochia Sipho 5mm. in thickness, m, medulla layer present in the cambium from
is
which on one or both sides the rows
fv, vascular bundle ; el, xylem ; cb, phloem ; /c
fascicular cambium ; \fc, interfascicnlar cam of cells take their origin. In many
bium ; p, phloem parenchyma ; pc, pericycle cases in which the cambial activity
sk, ring of sclerenchyma starch -sheath
e, ;
is confined to one side, according to
c, primary cortex cl, collencliyma in primary
;

cortex, (x 9.)
J. C. SCHOUTE, the original initial
cells are used up in the tissue forma-
tion, and new initial cells arise from adjoining cells of the ground-tissue.

Growth in Thickness of the Stem in Gymnosperms and Dicoty-

ledons. The cambium of the open vascular bundles of Gymnosperms
and Dicotyledons, which exhibit a growth in thickness, commences
its activity almost directly after the formation of the primary tissue.
The primary meristem remaining between the xylem and phloem of
the bundle continues its active growth as the cambium. Its cells are
full ofprotoplasm and continue to divide by means of tangential and
occasionally radial walls. The new cells thus continuously given oft'
from the initial cells toward the xylem and phloem sides of the bundles
experience another tangential division before attaining their definite
form as elements of the xylem or phloem portions. The vascular
bundles of Gymnosperms and Dicotyledons which undergo secondary
growth are usually arranged in a circle. After the cambium in the
bundles begins its activity, a zone of tangentially dividing tissue,
 MORPHOLOGY 1-29

called the INTERFASCJCULAK CAMBIUM, develops in the primary

medullary rays between the original bundles, and, uniting with the
cambium in the bundles, forms a complete cambium ring. This
cambium ring is thus composed of two distinct forms of meristematic
tissue for while the cambium of the bundles or the FASCICULAR
:

-.1C

rrp
FIG. 138. Transverse section of a stem of Aristolochia Siplw in tin; Hist year of its growth, showing
a vascular bundle with cambium in active division, p (Vascular .parenchyma i'lp proto-
xylein ; TO' and TO", vessels with bordered pits; ic, interfascicular cambium in continuation
with the fascicular cambium ; v, sieve-tubes cftp, protophloem pc, pericycle ; sk, inner part
;

of ring of sclerenchymatous fibres, (x 130.)

CAMBIUM consists of primary meristem

(p. 99), the connecting zone
of interfascicular cambium
of later development, and js consequently
is

a secondary meristem (p. 99). A

cross-section of a young stem of
Aristolochia Sipho, with the cambium ring in process of formation, is
represented in Fig. 137; in Fig. 138 a single bundle of the same
cross-section, more highly magnified, shows the fascicular cambium
with the interfascicular cambium to either side in a condition of
130 BOTANY

active division. \\7 ithin the

bundle may be seen two large vessels
(m"), in aincomplete state
still while in the adjoining primary
;

medullary rays the cells which give rise to the interfascicular

cambium may still be plainly distinguished. All the tissue arising
from the inner side of the cambium ring goes to form the secondary
WOOD, while that produced on the outside is termed secondary
BAST. The vascular portions of the wood form the WOOD STRANDS,
the sieve portions within the bast the BAST STRANDS. By the
activity of the interfascicular cambium, the primary medullary

Fir,. 139. Portion of a four-year-ol<l stem ot tin- 1'inc, 1'innxlsi/lcexti-i*, cut in winter. /, Trans-

verse view ; /, radial view ; t, tangential view ; /, spring wood ; s, autumn wood ; m, medulla ;

p, protoxylem ; 1, 2, 3, 4, the four successive annual rings of the wood ; i, junction of the wood
of successive years ma, ins', ?.<"', medullary rays in transverse, radial, and tangential view ;
;

ins", radial view of medullary rays in the bast ; c, cambium ring b, bust
; //. n-sin canals
; In;;

bark external to the first jwriderm layer, and formed from the primary cortex, (x H.)

rays are continued throughout both the wood and bast. As the
wood and bast strands enlarge, SECONDARY MEDULLARY RAYS are
developed from the fascicular cambium. In one direction the
secondary medullary rays terminate blindly in the wood, and in the
other in the bast the later they develop, the less deeply they
;

penetrate the tissues on either side of the cambium.

The primary medullary rays are therefore often distinguished as long, the
secondary as short medullary rays. The expression transvcrx- parenchyma is also
sometimes used to designate the medullary rays, winch arc in fact composed almost
exclusively of parenchymatous tissue. The cells given oil' by the initial layer of
the cambium for the formation of medullary ray- do not undergo a further tangential
division, as in other cases, but assume at once the character of medullary ray cells.
 MORPHOLOGY 131

The cambium have, for the most part, the shape of right-angled prisms, of
cells
which the radial diameter is smaller than the tangential. The ends of these prisms
are usually one-sided, tapering to a point, alternately on the right and left sides.
The length of the cambium cells varies in different plants, but those from which
mi'ilullary rays are formed are the
shorter. The primary vascular por-
tions of the bundles projecting into
the medulla constitute what is known
as the MEDULLARY SHEATH.

Owing to climatic varia-

tions, the cambial tissue of
woody plants exhibits a period-
ical activity which is expressed

by the formation of ANNUAL

KINGS of growth (Figs. 139,
141 A, 146). In spring, when
new shoots are being formed,
wider tracheal elements are
developed than in the follow-
ing seasons (Figs. 141 A, 148).
For this reason a difference is
perceptible between the EARLY
WOOD (spring wood), which is
composed of large elements
especially active in the convey-
ance of water (Fig. 141 /),
and the LATE WOOD (autumn
wood), consisting of narrow
elements which impart to a
stem its necessary rigidity
(Fig. 141 s}. Throughout the
greater part of the temperate
zone, the formation of wood
ceases in the latter part of
AugUSt Until the following ,, l(i uo ._ 1Magnml to illuslral( the secondary growth
.

-spring, when the larger ele- inthickneMCrftheitemaadvQofcofaGyBmoapenn

or Dicotyledon. Cambium, indicated by; dotted
mentS of the Spring WOod are , 11

, ! j f* . line ; m, pith ; 1, 2, 3, 4, successive animal rings

again developed. Owing to , jf W01K , ,, h lst
. . .
S( , % limit ,,, UW)I Imiin stem _,,,

the contrast in the struc- tap-r..t.

ture of the spring and the

autumn wood, the limits (Fig. 139 i)
between successive annual
rings of growth become so sharply defined as to be visible even
to the naked eye, and so serve as a means of computing the age
of a plant.

Under certain conditions tin- number of annual rings may exceed the number
of years of growth, MS. for instance, when MIDSUMMKII cii'wni occurs, such as
132 BOTANY PAKT I

commonly happens in the Oak, when, after the destruction of leaves by caterpillars,
a second formation of spring wood is occasioned by the new outgrowths thus
induced. In the wood of tropical plants the annual rings'may be entirely absent.
This occurs, for example, in the tropical Conifers of the genus Araucaria, which,
in this respect, show a marked contrast to the Conifers of the northern zone.
Any
interruption of growth, such as would occur during a drought, followed by a
period of renewed activity, may occasion the formation of annual rings even in
tropical plants.

Although a cessation in the formation of wood takes place so

Kio. 141. A, Transverse section of the wood of a Pineat'the junction of two annual linus. ./. Earl\
wood ; s, late wood ; t, bordered pit u, interposition of a new row of tracheides resulting from
;

the radial division of a cambium cell ; A, resin canals; m, medullary vays.; </, limit of lat>-
wood, (x 240.) If, Part of a transverse section of the stem of a Pine, s, Late wood; c,
cambium ; v, sieve-tubes ; p, bast parenchyma ; A-, cell.of bast parenchyma containing crystal :

cv, sieve-tubes, compressed and functionless ; HI, medullary ray. (x 240.)

early, the cambium tissue continues to form bast so long as climatic

conditions permit. As a rule, however, far'fewer elements are added
to the bast than to the wood. Owing to the continued growth in
length, the higher a cross-section is made of a gymnospermous or

dicotyledonous stem the fewer will be the annual rings. As the

diagram in Fig. 140 shows, the older rings disappear first on passing
towards the apex, and in the same way the older layers of the bast
are unrepresented nearer the apex. Up to a certain period, in the
age of woody plants, the elements of both wood and bast exhibit a
progressive increase in size.
The living elements may remain in a state of greater or less
 MORPHOLOGY 133

activity throughout the whole of the wood, extending even to the

pith ; such wood is called splint wood the Beech (Fagus sylvatica) :

may be quoted as an example. In other trees which form heart-wood,

the living elements die after a certain time, so that only dead tissues
ure found at a certain distance from the cambium. Before the death
of the living cells, they usually produce certain substances, such as
tannin and gums, which permeate the cell walls of the surrounding
elements, and also partially close their cavities. The tannins impart
to the dead wood a distinct colour, often very characteristic,
especially when it has been transformed into wood dyes, or so-called
XYLOCHROME. The tannin in the woody walls acts as a preservative
against decay, while the gums close the functionless water-courses of
the dead wood. The dead portion of the wood of a stem is called
the HEART-WOOD Or DURAMEN, in
contrast to the living SAP-WOOD
or ALBURNUM. Usually the splint
or sap wood is at once distinguish-
able from the heart-wood by its
lighter colour. In some stems,
however, the heart-wood does not
change colour.
its In that case,
as the protecting materials are
generally absent, it is liable to
decay, and then, as so often occurs
in the Willow, the stem becomes
hollow.
The sap-wood is limited, ac-
cording to the kind of wood, to
a larger or smaller number of the
younger annual rings, and tO it FIG. 142. Transverse section of a vessel fron
the heart- wood of Robinia Pscwlt'cttcia, closed
falls the task of water conduction.
by tyloses]; at a, a is shown the connection
'
between the tyloses and the cells from which
The distinction between sap- and
they have been formed, (x 300.)
heart- wood sharpest where the latter is
is

dark-coloured, as in the Oak, with its brown heart- wood, and in species of Diospyms,
whose black heart- wood furnishes. ebony. The darker the heart- wood, the harder
and more durable it usually is. The following may be mentioned as examples of
woods which yield dyes and colouring principles Haematoxylon campcchianum, L.
(Campeachy wood, logwood), with a red heart-wood from which H^MATOXYLIN is
extracted ; Pterocarpus santalinus, L. fil. (red sandal-wood), from the heart-wood
of which SANTALIN is obtained Caesalpinia brasiliensis, L., and G. ecltinata, Lam.
;

(Brazil wood, Pernambuco wood), with a red heart-wood which supplies BRASILIA ;

and the Alsage Orange, Madura aurantiaca, Nutt. (yellow Brazil wood), which has
a yellow heart- wood from which MORIN is derived. Inorganic substances may also
be deposited in the duramen thus calcium carbonate is found especially in the
;

vessels of the Elm and the 'Beech, while silicic acid occurs in those of the Teak
( Tectona grandis).
TYLOSES (Fig. 142) are also instrumental in closing the water-courses of the
134 BOTANY I-AHT i

cart -wood.
1 1 These are intrusive growths from living cells, which i>cnctr;itc tin-
cavities of the adjoining tracheal elements during the transition of the sap-wood
into heart-wood. In the formation of tyloses the closing membranes of the pits of
pitted vessels form bulging ingrowths into the cavities of the vessel. Such
ingrowths increase in size until several meet, and so more or less completely close
the cavities of the vessels into which they have intruded. The closing membrane
of the bordered pits in the heart- wood is pushed to one side, so that the torus presses
against the opening of the pit and completely closes it. According to H.
MA.YH ( 12B ), resin does not penetrate the walls of wood cells under normal con-
ditions ;
the wood of Conifers only becomes resinous through the impregnation
of the cell walls with resin, after they have become dried up through wounds or
other causes. The resin-ducts of Conifers may also be closed by the formation <>t
tyloses.

The elements of secondary growth differ in Gymnosperms and

Dicotyledons. The vascular strands of Gymnosperms are composed
almost exclusively of tracheides (Fig. 141 A}. These are provided
with bordered pits which are situated, for the most part, in their
radial walls. The tracheides of the spring-wood (/) have larger
cavities than those formed later (s). Parenchyma is also present in
the wood, though in relatively small amount ; in some Abietineae
resin-passages occur in it (Fig. 141 h).

Except in the Gnetaceae, true vessels are not found in the secondary growth,
nor in the primary vascular portions,, of the bundles of Gymnosperms. The wood
produced by the cambium consists of radial rows of tracheides, the number of
which is occasionally doubled by the radial division of a cambium cell (Fig.

141 A, a). The tracheides are often over a millimetre long, much longer than
the cambium cells from which they are developed. They attain this length by
a subsequent growth, during which their growing ends become pushed in between
one another. In addition to the tracheides, small amounts of wood parenchyma
are also produced in Gymnosperms by a transverse division of the cambium cells.
It is in the parenchymatous cell rows of the wood of Pines, Spruce-Firs, and
Larches that the schixogenous resin-ducts are produced (Fig. 141 A, h). In other
Conifers the wood parenchyma consists of simple rows of cells, which afterwards
become filled with resin.

Besides tracheides (t) and wood parenchyma (hp), other element >
take part in the composition of the secondary wood of a Dicotyledon ;

these are the vessels (tracheae, g), and the wood fibres (h) (Fig.
145 A, B}. The cells of the wood parenchyma are short and have
abundant contents, the woods fibres are thick- walled, long cells with
pointed ends. The elements with wider lumens, especially the
vessels, are abundant in the spring-wood, in which water conduction
is important. The autumn -wood, on the other hand, consists of
narrow elements, among which the wood fibres, which contribute to
the rigidity of the plant, are numerous. On account of these
differences between spring and autumn wood the annual rings are
well marked (Fig. 148).
SECT. I MORPHOLOGY 135

All the elements entering into the formation of the wood of Dicotyledons can be
derived from the two classes of tissue already met with in the Gymnosperms, the
tracheal tissue and the parenchymatous tissue of the wood. The tracheides and
vessels belong to the former class, while under the parenchymatous tissue are
included the wood parenchyma, fibrous cells of greater length but with similar
contents (Fig. 145 ef), and the wood fibres.
The tracheal tissue consists of elements which lose their living contents at an
early stage, and in their fully developed condition are in reality only dead cell
cavities. In this class are included TRACHEIDES having relatively wide lumina and
large bordered pits, and ultimately also spirally thickened walls, which serve as water-

sni

FIG. 143. Radial section of a Pine stem, at the junction of the wood and bast, s, Late tracheides ;
t, bordered pits ; c, cambium ; v, sieve-tubes ; at, sieve-pits ; tin, tracheidal medullary raj-
cells sm, medullary ray cells in the wood, containing starch ; sm', the same, in the, bast ;' em,
;

medullary ray cells, with albuminous contents, (x 240.)

A, t) ; VASCULAR TRACHEIDES (gt), with similar functions, but with

carriers (Fig. 145
the structure and thickening of vessels FIBRE TRACHKIDES (ft), with small lumina
;

and pointed ends, having only small, obliquely elongated bordered pits, and, in ex-
treme cases, exercising merely mechanical functions and finally TRACHEA (</),
;

formed by cell fusion, and provided with all the different forms of thickenings by
which they are distinguished as annular, spiral, reticulate, or pitted vessels. All
vessels function as water-carriers. If they have small lumina and resemble
tracheides, they may be distinguished as TRACHEIDAL VESSELS (tg) ; if, as is
generally the case, they, have bordered pits on their lateral walls, they
are usually provided with tertiary thickening layers in the. form of thin,
spiral bands (Fig. 149 m). In the parenchymatous tissue of the wood the
145 B) generally retain their living contents, and never develop the true
cells (Fig.
bordered pits with a torus in the closing membrane, which are so characteristic
136 BOTANY PART I

of the water-conducting elements. All tissues of this class may be best derived
from wood parenchyma. The wood parenchyma
produced by transverse is
divisions of the cambium cells, and accordingly consists of rows of cells (hp] with
transverse division walls, and others obliquely disposed, which correspond to the
alternately differently pointed ends of the cambium mother cells. The cells of the
wood parenchyma are provided with simple round or elliptical pits, varying in size in
different kinds of wood they generally contain starch, and some of them also take
;

up bye-products, resulting from metabolism, or

from the chemical changes taking. place within
A plant in the processes of its nutrition and
growth. The cells having the closest resem-
blance to those of typical wood parenchyma
are the so-called FIBROUS CELLS (ef). In their
\-t
contents, as well as in their wall thickenings,
they are similar to the cells of wood parenchyma,
but each [is formed directly from one entire
cambium cell. In' their formation, the cells of
the cambium tissue become more or less elong-
ated and fibrous. The LIBRIFORM FIBRES or
WOOD FIBRES (h) have a similar origin, but are
even more elongated and have thicker walls,
and, at the same time, narrow, obliquely elong-
ated, simple pits. Although the wood fibres

,v may continue living, in the more extremely

developed forms (h) they lose their living
contents. They are then filled with air, and
their function -merely mechanical.
is Under
certain conditions, by later transverse divisions,
the libriform fibres may become transformed
into SEPTATE WOOD FIBRES (gh). The trans-
verse septa thus formed remain thin, and form
a striking contrast to the more strongly thick-
ened lateral walls. While the tracheal tissues
\

FIG. 144. Tangential section <>t the

are engaged in providing for the conduction of
autunm-wood of a Pine, t, Bordered water, the duty of conducting and storing the
pit; tm, tracheidal medullary ray products of assimilation, in particular the
cells; am, medullary ray cells con- performed by the parenchy-
is
' carbohydrates,
matous tissues of the wood. Both forms of
on one side; i, intercellular SJKII-I- in
the medullary ray. (X 240.) however, aid in maintaining the rigidity
tissue,
of the plant body, and, in their most extreme
development, furnish such elements as the fibre tracheides on the one hand, and
on the other the empty wood fibres, which are only capable of performing
mechanical functions.
The wood of Dicotyledons is made up of the elements of these two classes of
tissue, the tracheal and the parenchymatous, but all the different elements are not
necessarily represented inany one kind of wood.
Drimys, a genus belonging to the Magnliaceae and two genera of the related
order Trochodeiidraceae, are the only Dicotyledons the wood of which is formed of
tracheides only. These Dicotyledons closely resemble the Conifers in structure.
In numerous Leguminosae, Willows, Poplars, and species of Ficus, on the other
hand, the tracheal tissues are only represented by vessels, which perform the task
 MORPHOLOGY 137

of water conduction. In the wood strands of these plants there are also present
wood parenchyma and a large proportion of wood fibres. The vessels in climbing
plants (lianas) are especially wide.
The distribution of the living elements in the wood strands always bears a dis-
tinct relation to the water-courses which they accompany,
enclosing them in a more
or less complete sheath. The living cells adjoining the tracheal elements are in
communication with them by means of one-sided bordered pits. When such pits

ft

Fio. 145. .4, Elements of the tracheal tissue of the wood

; diagrammatic. Elements of the
/.',

parenchymatous tissue of the wood ; diagrammatic. For description see text.

occur between living cells and tracheal (elements the pit cavities are absent on the
side of the living cell, but present in the tracheal elements they differ from the
;

true bordered pits in the absence of a torus on the pit membrane.

The Elements of the Secondary Phloem in Gymnosperms and

woody Dicotyledons are sieve-tubes, or these together with companion
cells, bast parenchyma with abundant
cell contents, and long narrow

bast fibres with strongly thickened walls. The sieve-tubes serve to

conduct proteid materials ; the companion cells, or in their absence
special rows of the bast parenchyma,
take up substances from the
138 BOTANY I'AUT I

sieve-tubes storage and conduction of carbohydrates take place in

;

the parenchyma in which bye-products of metabolism, such as tannins

and calcium oxalate, also accumulate.
As in the case of the wood, the elements of the bast may be referred to two
forms of tissue, the sieve-tube and the parenchymatous. The former is represented
by the sieve-tubes or by these together with companion cells, the parenchymatous
portion by the phloem parenchyma and the bast fibres, between which there are
intermediate forms of element.

In the bast strands of Gymnosperms, the phloem elements pro-

duced by the cambium (Fig. 141 ft, c) consist solely of sieve-tubes,

-.".

;'//
fZfe.~-*J tm

;
.T
- -

-pm

FIG. 146. Transverse section of a stem of Tilin KM.. 147.- A radial section of the wool of
in the fourth year of its growth. Tiliii ulmijuliii, showing a small medul-
pr, Primary cortex ; c, cambium ring ; cr, bast ; lary ray. g, Vessel ; I, wood fibres ; ////.

pm, primary medullary ray; pm', expanded ex- medullary ray cells in communication
tremity of a primary medullary ray am, second-
; with the water -courses by means of
ary medullary ray ; g, limit of third year's wood. pits snt, conducting cells of the medul-
;

(x 6.) lary ray. (x '240.)

the parenchymatous cells of the bast parenchyma (p and /,), and, in

certain cases, of bast fibres. These elements of the bast generally
form alternating bands.
The Araucarieae, Taxineae, and Cupressineae have definite, vertical rows of
bast parenchyma cells which take the place of the companion cells wanting in
these plants. At a certain distance from the cambium the sieve-plates become
overlaid by callus. During the vegetative period following their development,
the sieve-tubes become empty and compressed together (Fig. 141 B, cv). The
rows of bast parenchyma cells containing albuminous substances, which are
found in some Conifers, undergo disorganisation at the same time as the adjacent
sieve-tubes the bast parenchyma cells which contain starch, on the other hand,
;

continue living for years, and even increase in size, while the sieve-tubes become
compressed.

The elements of the phloem tissue included in the bast strands

SECT. I MORPHOLOGY 139

of woody Dicotyledons (Fig. 150) are represented by SIEVE-TUBES (v)

and COMPANION CELLS (c). To the parenchymatous tissues of the
bast belong BAST PARENCHYMA (p), BAST FIBRES (/) and transitional
forms between them.

The bast fibres,' like the fibres of the wood, may occur in an unthickened' form
as FIBROUS CELLS, either with or without living contents, or they may be filled
with starch, or finally may become septate.
The elements of the
bast of Dicotyledons fre-
quently exhibit a great
regularity in their ar-

rangement. Thus in the

Linn- (Fig. 150) there
is a regular alternation
of sieve -tubes (v) and

companion cells (c), bast

parenchyma containing
starch (p) and crystals
(Jc), tangentially flat-

tened bast parenchyma

(p), and then another
zone of sieve-tubes. The
sievo-tubes of Dicotyle-
dons as of Gymnosperms
only remain functional
for a short time, after
which they become
empty and crushed. The
companion cells, which
are sister cells of the
sieve-tubes, undergo the yiff
^ f
same fate. The starch-
containing parenchyma, !'[<;. 148. Portion of ;i transverse section of the wood of 'Cilia
on the other hand, may idmifolia. TO, Large pitted vessel t, tracheides
; I, ; wood
remain for years un- fibre ; p, v.ood parenchyma r, medullary ray. ( x 540.)
;

altered. The differences

in the appearance of the bast of dicotyledonous trees are due to the greater or less
diameter of' the sieve-tubes, the presence or absence, of bast' fibres, and the arrange-
ment of the various elements.

The Medullary Rays of the Gymnosperms (Fig. 139 ms) and

woody Dicotyledons (Fig. 146 pm, sm) form radial bands, composed
wholly or in part of parenchymatous elements. Their function is to
supply the cambium and wood with the products formed in the leaves
and conveyed away by the bast they also conduct water outwards ;

from the xylem. The medullary rays in this way link together by
radial bands of living cells the protoplasm-containing elements of the
bast and wood, thus uniting all the separate living tissues of the stem.
The medullary rays are in turn accompanied or, if many-layered,
140 BOTAXY PART I

traversed by intercellular spaces filled with air. These, beginning in

the periphery of the stem, penetrate the cambium and communicate
with all the intercellular spaces throughout the living elements of the
wood and bast. All the living elements are kept in communication
with the atmosphere by means of the
spaces of the medullary
intercellular
rays, and the necessary interchange of
gases is thus rendered possible.
The substances contained in the parts of
the medullary rays within the wood, chiefly
consisting of starch, tannins, resin, and crystals,
are essentially the same as those in the woo. 1

parenchyma. In the medullary rays of certain

Gymnosperms, particularly in the Pine, single
rows of without living contents and situ-
cells,
ated usually at the margin of the medullary
bands, become tracheidal in structure (Figs. 143,
144 (m), and united with one another and with
the tracheides by means of bordered pits. Their
purpose is to facilitate the transfer of water
radially between the tracheides. In other
Conifers, where such tracheidal elements are
not found in the medullary rays, bordered pits
are developed in the tangential walls of the
tracheides of the autumn- wood, and by means of
them the transfer of water in a radial direction
is effected. The living cells of the medullary
rays of the wood bear the same relation to the
water-carriers as does the wood parenchyma,
like them are connected with the water-
and
conducting elements by means of bordered pits
on one side of the cell wall. They take up water
from them and give it out again, as it may be
FIG. 149. Tangential section of the wood needed, toother
living cells on the other hand, ;

of Till,, ..tmifoli,,. m. Pitted vessel ;

in the wring, at the beginning of the season
traoheides wood paren-
t, spiral

,-hyma; wood llJ.n-s

I,
j>,
;

; r,
of gH"**, the Jr press
m.Nlullary
* m
.

* he water-cour.ses

rays, (x 160.) the products of assimilation, in particular

glucose and small quantities of albuminates,
in order that these substances may be transferred in the quickest way to the points
of consumption. Accordingly, during the winter and in the beginning of spring,
sugar and albumen may be detected in the tracheal elements ( ), and may thm
}26

"
be obtained from the watery sap of "bleeding trees, or from artificial borings or
incisions, particularly in such trees as the Maple, Birch, and Hornbeam. In the
wood of Dicotyledons it is usually only special rows of the medullary ray cells
which stand in such close relation with the tracheal tissues. In these special rows,
generally on the margins of the medullary rays, the cells- are elongated vertically,
and on that account have been distinguished as VERTICAL MEMUI.LAUY KAY CELLS.
Theiother cells, or those of the middle layers of the medullary bauds, on the other
hand, are called HORIZONTAL MEDULLARY KAY CELLS they are narrower and
;

I27
more elongated radially ( ). These have no especial connection with the tracheal
SECT. I MORPHOLOGY 141

elements, but are designed for conducting and storing assimilated substances.
Within the bast xone the medullary rays of Dicotyledons have a simpler structure
than in the wood. It is evident, not only from the pits between the cells of the
medullary rays and the bast parenchyma, but also from the similar relations
in Dicotyledons between the medullary ray cells and the companion cells of the
sieve-tubes, that the function of the cells of the medullary rays in the phloem
is up substances passing down the bast strands.
to take
In the Pine and other Abietineae, whose bast parenchyma is devoid of cells
functioning as conductors of albuminous matter, their place is taken in this respect
by rows of medullary ray cells (Fig. 143 em}. These maintain an intimate connec-
tion with the sieve-tubes by means of sieve-pits. They lose their contents in the

Fio. 150. Portion of a transverse section of the bast of Tilin ulmifnli". v. Sieve-tubes ; v*, sieve-
plate c, companion cells; k, cells of bast parenchyma containing crystals
; ;
;>, bast pui-en-

chyma ; f. bast fibres /, medullary ray. (x :>W.)

;

same manner as the sieve-tubes, 'and, like them, become compressed and dis-

organised. On
the other hand, the cells of the medullary rays, which contain
starch, like the similar cells of the bast parenchyma, increase in size, and continue
living for years.
The width and height of the medullary rays may be more easily determined
from tangential than from radial sections. In such tangential sections the medul-
lary rays appear spindle-shaped (Figs. 144, 119). With few exceptions, as in the
Oak and Beech, the medullary rays are of relatively small size. The Oak, in
addition to numerous small medullary rays, has other larger rays which may be as
much as a millimetre broad and a decimetre high. In the Poplar, Willow, and
Box the medullary rays are so extremely small that they are scarcely visible, even
with the aid of a magnifying glass. The height of the broad primary rays of many
lianes, on the other hand, may be equal to that of a whole internode. In certain
Conifers, resin-ducts occur not only in the wood, but also in the broader medullary
142 BOTANY

rays. These radial resin-ducts are in communication with the vertical ducts. It
is due amount of resin exudes from wounds in Pine or
to this fact that such a large
Fir trees.

Secondary Thickening of the Root in Gymnosperms and

Dicotyledons. The ROOTS of Gymnosperms and Dicotyledons, in
which the stems increase in thickness, show a similar GROWTH IN
THICKNESS (
12S
).
When secondary
growth begins in a root with its
xylem and phloem strands alter-
nating with each other (Figs. 1 24,

131), layers of cambium arise on

the inside of the phloem strands,
through the division of the funda-
mental tissue these give off wood
:

elements towards the centre of the

root, and bast towards the periphery.
These cambium layers soon meet in
the pericycle, just outside the xylem
strands, and so form a complete
cambial ring. In Fig. 151 A, this
process is diagrammatically repre-
sented. As a result of the activity
of its cells the cambium ring soon
loses its sinuous form, and becomes
circular. In front of the primary
vascular strands (g'),
the cambium
produces medullary ray tissue, and
this constitutes the broadest medul-

lary rays which lead inwards to the

strands of primary xylem (Fig. 151
Fio. 151. Diagrammatic representation of the B). A crOSS-SCCtion of SUch a rOOt,
rowth in thickness of a dicotyledonous
in which the secO iidary growth has
root, pr, Primary cortex f,
; cambium
ring; g ; primary vascular strand; continued for sonieyears,can scarcely
,

primary phloem strand; j, pericycle; c, De distinguished from a Cl'OSS-section

of a stem. By careful examina-

tion, however, the presence of prim-
ary xylem in the centre of the root can be discovered, and its nature
thus determined. The wood of the root is also more porous than
in the stem, and bears a close resemblance to spring-wood. On
account of this lack of differentiation in the wood, the annual rings
of growth are less distinctly defined in roots than in steins.
In the root and its branches, as in the stem, the annual zones of
secondary growth become less numerous on approaching the growing
points (Fig. 140).
Extraordinary <lc\ i;itii>ns from
1U9
Anomalous forms of Growth in Thickness ( 1.

the usual type of secondary ^rowi li arc aH'onli-d l>y

and routs <>f Jyiiino-
-i>mr .-inns (
 MORPHOLOGY 143

sperms and Dicotyledons. Among the Gymnosperms in the Cycadaceae and

certain species of Gnetum, in tlie Chenopodiaceae, Amarantaceae, Nyctaginaceae,
Phytolaccaceae, and other families of Dicotyledons, the cambium which has been
formed in the ordinary manner soon loses its function, and a new cambium ring is
developed external to the bast zone, for the most part in the pericycle, or in a
tissue derived from it. This cambium ring forms wood on the inside and bast on
the outside, with the accompanying medullary rays. It then ceases to divide, and
a new ring takes its place. This process is repeated, and ultimately leads to the
formation of concentric rings of wood and bast, which, in cross-sections of the sugar-
beet, may be dist nguished with the naked eye. These concentric zones may be
stillmore plainly seen in a cross-section of Mucuna altissima (Fig. 152), a liane
belonging to the order Papilionaceae. The stem shows in this case an inner axis
of wood (1) suirounded by a zone of bast (1*) next follows a cylinder of wood (2)
;

and bast (2*), and finally a third (3, 3*) in process of formation in the pericycle.

Ki<;. Iji'. --Tr.-insviTsi' section of the stem of Mitcuiui altiasima. 1, -2, 'A.
Successively formed
xi nit's of wood ; 1*, 2*, 3*, successively formed zones of bast. (] nat. si/p.)

An extraordinary appearance is exhibited by cross-sections of stems, which show

several separate wood cylinders (Fig. 153). Such a structure is peculiar to various
tropical liancs of the genera Serjania and Paullinia belonging to the family
Sapindaceae. This anomalous condition arises from the unusual position of the
primary vascular bundles, which are not arranged in a circle but form a deeply
lobed ring so that, by the development of inter fascicular cambium, the cambium
;

of each lobe is united into a separate cambium ring. Each of these rings, in-
dependently of the others, then gives rise to wood and bast (Fig. 153). A very
peculiar structure is exhibited by many lianes of the Bignoniaceae, the wood of
which is cleft by radially projecting masses of bast (Fig. 154). The primary stem
of the Bignoniaceae shows the ordinary circular arrangement of the vascular
bundles. Wood and bast are at first produced from the cambium ring in the usual
manner, and an inner, normal wood cylinder of AXIAL WOOD is formed. Such normally
formed axial wood cylinders are common to many, otherwise abnormally developed
lianes. The cambium ring of the Bignoniaceae, after performing for a time its
normal functions, begins, at certain points, to give off internally only a very small
quantity of wood, and externally a correspondingly large amount of bast. As a
result of this, deep wedges of irregularly widening bast project into the outer
144 BOTANY I'ART I

so-called PERIAXIAL WOOD (Fig. 154). The originally complete cambium becomes
thereby broken into longitudinal bands, which are broader in front of the projecting
wood than at the apices of the bast wedges. As the periaxial wood is always
developed from the inside, and the wedges of bast from the outside of their re-
spective cambium bands, they extend past each other without forming any lateral
connection.
The Formation of Knots. The knots or streaks which add so greatly to the
technical value of certain woods depend on an unusually bent or interwoven course
of the elements of the wood. Their origin is due to the stimulus of wounding,

Fin. 153. Transvrise section ol

the stem of Serjania Lnniot-
tenna. sk, Part of the ru]>-
tured .si-lerenchymatous rinj;
of the pericycle ; I and I*, bast
/ones ; Ig, wood ; m, medulla.
<x2.)
}'i>:. 154. Transverse section of the stem of one <>t

the Bignoniaceae. (Nat. size.)

to the effects of parasites, the pressure exerted by lateral branches which are in-
creasing in thickness, or to altered cambial activity. Larger knots are produced
by the origin of numerous adventitious buds, especially after wounding a finer ;

marking by the widening of the medullary rays, which may then appear circular
in tangential section, and influence the course of the adjacent elements of the
13
xylem ( ).

Secondary Growth of Monocotyledons. There are certain morio-

cotyledonous plant families and genera, especially Dracaena, Yucca,
Aloe, and the Dioscoreaceae, in the stems and roots of which a cambium
ring is developed. As in such cases, the cambium ring generally
arises in the pericycle, outside the scattered vascular bundles and
from the fundamental tissue, it is a secondary meristem ; it does not,
as in Dicotyledons and Gymnosperms, produce continuously wood
and bast in opposite directions, but, instead, only new tissue to the
inner side. This later becomes differentiated into closed vascular
bundles embedded in a thick-walled parenchymatous ground tissue.
IS1
The Gambia 1 ring in these cases is only active to one side ( ), and its initial
 MORPHOLOGY 145

cells are replaced from the cells of the ground tissue on the outer side of the
ring (p. 128).

At a later period the cambial ring commences to form cells to the

outer side which increase the thickness of the cortex as a thin-walled,
parenchy matous ground
tissue (Fig. 155). When the & r

^ /
\> ^ '-' -

cambium has thus become

activeon both sides itpresents -
;~4=
the appearance of a single
initial layer.
As shownin the diagram

(Fig. 156)the growth in

thickness of a Monocotyledon
differs essentially from that of
a Grymnosperm or Dicotyledon
-
represented diagrammatic
ally in Fig. 140. The stem
of a Dracaena on which the
diagram is based is represented
as unbranched. Its primary
central cylinder (pc) increases
in diameter as the stem grows
in length, and thus forms an
inverted cone. It is surrounded
by the zones of secondary
growth, the amount of which
diminishes on passing towards
the growing point. The
secondary growth of the stem
is continuous with that of the

roots.
Secondary Growth of
Leaves ( 132 ) is always very
slight, and is confined to a
few Coniferae and Dicotyle-
FIG. 155.Transverse section of the stein of: Cordyline
dons with evergreen leaves. (Dracaena) rubra. /', Primary vascular bundles;
The increase in thickness is /", secondary vascular bundles: /'", leaf -trace
bundle within the primary cortex 'm, parenchy- ;

due to the presence of a tnatous fundamental tissue; bund le- sheath

s, ; *,

fascicular cambium. This tracheides ; c, cambium ring ; cr, cortex, the outer

portion being primary, the inner secondary cortex ;

produces little or no xylem,
ph, cork cambium ; I, cork ; r, bundles of raphides.
thenew elements beingmostly (x 30.)
added to the phloem. Any
elements formed on the side next the xylem are water-conducting,
while those adjoining the phloem mostly become sieve-tubes.
Periderm ( 133 ). It is very seldom that the epidermis, by the
division of its own cells, is in a condition to keep pace for any length
L
 HOT AN V

of time with the increasing dimensions of the stem. This, however,

is the case with the Mistletoe (Viscum album), the number of epidermal

cells of which is continually augmented by the formation of new

lateral walls, and new cuticular layers are formed where the old
cuticle becomes ruptured. On older portions of the stem, however,
thickening layers form in single epidermal cells or groups of
epidermal cells, and at the same spots similarly thickened cortical
cells are developed. The epidermal cells thus cut off die, and
the original epidermis is more or less completely replaced on such
old regions of the stem. In the
great majority of plants in which
the epidermal layer persists for
years it is able to meet the
demands made upon it by the
growth in thickness of the stem
without the assistance of the under-
lying cortex. The stems of one of
the Maples (Acer striatuw), even
when a foot or more in thickness
and over forty years old, remain
covered with a living, growing,
epidermal layer. The striations to
which the specific name of this
plant refers are caused by a white
wax exuding from splits in the
outer walls. As a rufe, however,
the epidermis on stems which grow
, in thickness becomes stretched and
FIG. 15i>. Diagram of the secondary growth of
' ., ,
f^. . .

the stem and root of Dracoew. pc, Central finally ruptured. The tlSSUC of
cylinder; s, secondary tissue; c, cambium the primary COI'tCX and of the bast
indicated by a dotte-l line;,, .root. (Adapted
fe th expansion am j division of its
from SCHOI-TK.) ,
cells, can accommodate itself more
easily than the epidermis to the
increased dimensions of the stem,

arising from the growth in thickness of the central cylinder. This

noticeable in the primary
process of accommodation is particularly
medullary rays (Fig. 146, pm) between the primary phloem. The
formation of the PERIDERM generally begins during the first vegetative
period,after the secondary growth has reached a certain stage. The
commencement of its formation is indicated by the brown colour of
the external surface of the stem, which remains green so long as the
epidermis continues alive. The formation of periderm usually occurs
at a later period in those plants which have a persisting epidermis ;
it is practically wanting only in the species of Mistletoe. The periderm
is derived from a secondary meristem, termed the CORK CAMBIUM or

I'HKLLOGEN. This phellogen may arise, in the epidermis, in a deeper

layer of cells of the primary cortex, or even in the pericycle itself.
 MORPHOLOGY 147

124
The phellogen ( )
is as a rule a cambium with a persisting initial
layer from which cork-cells are produced to the outside and phello-
derm to the inside.

The productionof cork -cells is usually greater than that of phelloderm. If the

phelloderm forms only cork-cells the initial cells are often used up in the process,
and are then replaced from the adjoining ground tissue (p. 128). This is found
among Monocotyledons and also in some Dicotyledons, e.g. Valcriuna ojficinalis.

The young cork-cells usually have a tabular shape, fit closely

together without intercellular spaces, and possess suberised, secondary,
thickening layers. The cork cells usually possess brown walls.

~cof

Fin. 157. Transverse section of the outer part of :i one-year-ol'l twi^ of 1'ynis communis made in
autumn. It shows the commencement of the formation of the periderm. />, cork ; pg,
phellogen ; pd, phelloderm ; ml, collenchyma. The cork-cells have their outer walls thickened
and have brown dead contents, (x 500.)

They may be thin (Fig. 155 /) or thick, frequently thickened on one

side (Fig. 157), and occasionally to such an extent that they are
known as stone Thin-walled cork-cells usually contain air ;
cork.
those with thickened frequently have reddish brown cell
walls -

contents (phlobaphene) consisting mainly of tannins and products

derived from them. The cork tissue frequently shows an alternation
of narrow thick-walled with thin-walled and larger cells. These
layers mark annual growths. Even thin corky walls, being imperme-
able to water, prevent the loss of moisture by transpiration. Thicker
layers of cork afford mechanical protection and hinder the invasion
of the plant by parasites.
The phelloderm formed to the inner side of the phellogen in many
cases, such as the Currant, attains a considerable thickness. Its
cells retain their living protoplasm, and usually contain chloroplasts.
148 BOTANY PART i

They ultimately become rounded off, so that intercellular spaces are

formed between them and serve to increase the thickness of the living
cortex. The term periderm includes both cork and phelloderm.

The cork of the Cork-oak >//or//.v .Sti/n :/) is formed of broad

'
t layers of .soft
large alternating with narrow layers of thinner cells, which mark the limit
cells,
of the annual growth. This may be seen in bottle-corks. The first, spon-
taneously developed cork of the Cork-oak is stripped off, whereupon a new phellogen
is formed in the deeper-lying tissue. The cork thus produced is removed every six
or eight years, and furnishes the cork of economic value.
In many cases the phellogen takes its origin in the epidermis (Fig. 157 py).
This is the case in the Willow, in all Pomaceae, and in a number of other woody
plants. The epidermal cells become divided into outer and inner cells, the
latter ofwhich assume the function of a phellogen. More frequently the phello-
gen develops from the layer of cells next adjoining the epidermis, as, for example,
in the Elder (Sambucus nigra), where it takes its origin from the outermost layer
of collenchyma (Fig. 158, ph). The epidermis in this case covers the developing

layer of cork.
True cork is confined to Ophioylossiuu among the Cryptogams. The fleshy
stipules of the Marattiaceae are covered with a brown layer arising by the irregular
division of sub-epidermal cells; the walls of this tissue are humified and it may
134
be termed pseudo-periderm ( ).

Bark.All tissues external to the phellogen are cut off from food
supplies, and consequently die. When the first cork layer has its
origin deep within the stem, a BAKK of considerable thickness is formed
through the ensuing death of the peripheral tissues. If the cork layer
formed by the phellogen be thin, the stem has a smooth surface, as
in the Beech if it produces thicker cork layers, the surface of the
;

stem appears rough and full of fissures, as is the case in the Cork-oak.
The primary phellogen generally ceases its activity after a short time,
and another deeper-lying phellogen is formed. After a time this
new phellogen discontinues its functions, and another (Fig. 159) is
developed, as in the case of Quercus sessiliflora, until ultimately the
phellogen comes to be formed in secondary bast parenchyma instead
of in the primary tissue. That portion of the bast cut off' by the
periderm loses its nutritive contents and only retains waste products.
If the layers of the secondary periderm constitute only arcs of the
stem circumference, the bark will be thrown off in scales, as in the
SCALY BARK of the Pine and Plane tree ; if, on the contrary, the
periderm layers form complete concentric rings, then hollow cylinders
of the cortical tissues are transformed into the so-called RINGED BARK,
such as is found in the Grape-vine, Cherry, Clematis, and Honeysuckle.
Bark which is not easily detached becomes cracked by the continued
growth in thickness of the stem, and has then the furrowed appear-
ance so characteristic of the majority of old tree-trunks. The usual
brown or red colour of bark, just as in similarly coloured heart-wood,
is occasioned by the presence of tannins, to the preservative qualities
 MORPHOLOGY 149

of which is due the great resistance of bark to the action of

destructive agencies. The peculiar white colour of Birch-bark is
caused by the presence of betulin (birch-resin) in the cells.
In roots which grow in thickness the phellogen usually develops
in the pericycle (Fig. 151 B, k), and in consequence of this the

primary cortex of the roots dies and peels off. The succeeding
phellogen layers are formed in exactly the same way in the root as
in the stem.
13f>
Lenticels ).
In most woody plants, particularly in Dicotyle-
(

dons, cortical pores, or LENTICELS (Fig. 158), make their appearance

simultaneously with the formation of periderm. The lenticels take
their origin in a phellogen layer (pi) which, in the case of peripheral

FIG. 158. Transverse section of a lent.icel of Sambucvs nigra. . Epidermis ; /ih, phellogen ;

/, complementary d-lls ; /il, phellogen of the lenticel ; pd, phello<lerni. (x 90.)

cork formation, almost always develops directly under the stomata.

The phellogen, from which the lenticels arise, unlike the cork phellogen,
does not form cork cells, but a lenticel tissue composed of COMPLE-
MENTARY CELLS (/) with intercellular spaces between them. On the
inside, however, a phelloderm is regularly derived from the phellogen.
The complementary cells press the epidermis outwards and finally
rupture it. Where the complementary cells are only loosely united,
intermediate bands or closing layers are developed from the phellogen
alternately with the layers of looser tissue the closing layers become ;

eventually ruptured. The complementary cells are as a rule not

suberised, those of the closing bands are corky and lignified. The
cork-forming phellogen joins the phellogen of the lenticels at its
margins. In cases where the cork is more deeply seated in the inner
tissue, the lenticels begin their development at a corresponding
depth from the layer of phellogen. Lenticels serve to maintain the
150 1JOTANY

gaseous interchange between the outside air and the gases present in
the intercellular spaces of the plant.

Certain aerial roots such as those of the Orchids and the respiratory roots of
>\v.-iinp plants possess lenticels the
structure of which deviates more or
lessfrom the usual type ; these are
termed piieumathodcs.

Absciss The fall

Layers.
13
of the leaves ofwoody plants ( )
is
brought about by means of a
layer of parenchymatous tissue
which is formed at the base of
the leaf -stalk shortly before
the leaf -fall. The cells of this
tissue separate from one another

owing to their middle lamellae

becoming mucilaginous. With-
in this absciss layer all prosen-

chymatous elements of the

petiole are greatly reduced ;

only the tracheal elements are

lignified, and these are broken
through when the leaf separates.
The protection of the leaf-scar
is effected in the first place by
the lignification and suberisa-
tion of the walls of the exposed
cells, and later by the formation
of a layer of cork continuous
with the periderm covering the
stem. The ends of the tracheal
Fio. Transverse section of the
1')!'. peripheral elements in the scar become
tissues of the stem of Quercus xessiliflora. 1, 2, 3,

Successively formed layers of cork ; pr, primary

occluded by wound -gum or
cortex, modified by subsequent growth ; in- thylosis Or by both ; the ends
ternally to ,* 1-ricycle;*c, sclerenchymatous
f fcfe gieve . tubes become COm-
fibres, from the ruptured ring of sclerenchy- .

matous fibres of the pericycle ; s, subsequently pressed and The

lignified.
formed sclereids; s', sclereids, of secondary leaflets of leaves
bast fibres with accompanying crystal
Compound
growth ; cr, 11 r j e .

ceils; I-, ceils, with aggregate crystals. AII the usually become separated from
tissue external to the innermost layer of cork the rhachis by absciss layers,
is dead and discoloured, and has become trans-
Herbaceous plants exhibit as a
formed into lark, (x 22'..) , , - \ . . ,
n ,.,..
rule no definite leaf-fall ( 137 ).
13S
Healing* of ( ).
Wounds
Lost portions of tissue are in the lower
plants frequently regenerated directly, while this only occurs to a
13
very limited extent in more highly organised plants ( ). In the
latter regeneration is limited to the periderm, and in a few instances
 MORPHOLOGY 151

to parts of the vascular bundles. In the simplest cases among

land plants the wounded cells die and become brown and dry,
while the walls of the underlying uninjured cells become lignified.
The protection of wounded surfaces takes place in this way in the
Cryptogams and when the wound is very small in Phanerogams.
In the case of larger wounds in the latter a cork-cambium, forming
wound-cork, develops below the lignified cells. This cork is lignified
as well as suberised ( 14 ). Suberisation of the walls of the cells
immediately underlying the wound may precede the formation of
cork. This occurs in seed potatoes which are divided up before
planting and left one or two days exposed to the air. On this
141
depends their resistance to injurious influences in the soil ( ).
Secretory receptacles in wounds become closed by thyloses. Latici-
ferous elements are first closed by the coagulation of the latex at
the point of wounding and below this a cell-wall is formed ( 142 ).
In woody plants a so-called CALLUS is formed by the active
growth of the living cells bordering on the wound. These swollen
growths close together over the Avound, and by the suberisa-
tion of their cell-walls provide a sufficient protection. Generally,
however, a cork-forming phellogen arises in the periphery of the
callus. In stems of Gymnosperms and Dicotyledons, wounds which
extend into the wood become surrounded and finally overcapped by
an outgrowth of tissue arising from the exposed cambium. While
the callus tissue is still in
process' of gradually overgrowing the
wounded surface,an outer protective covering of cork is developed ;
at the same time a new cambium is formed within the callus,
through a differentiation of an inner layer of cells, continuous
with the cambium of the stem. When the margins of the over-
growing callus tissue ultimately meet and close together over the
wound, the edges of its cambium unite and form a complete cambial
layer, continuing the cambium of the stem over the surface of the
wound. The wood formed by this new cambium never coalesces
with the old wood. Accordingly, marks cut deep enough to penetrate
the wood are merely covered over by the new wood, and may
afterwards be found within the stem. In like manner, the ends of
severed branches may in time become so completely overgrown as to
be concealed from view. The growing points of adventitious shoots
often arise in such masses of callus. As the wood produced over
wounds differs in structure from normal wood, it has been distin-
guished as CALLUS WOOD. It consists at first of almost isodiametrical

cells, which are, however, eventually followed by more elongated cell

forms. At the base of cut-off portions of plants which are used as
cuttings a callus formation which according to the kind of plant is
more or less active, takes place. In extreme cases the callus forms pro-
jecting masses within which the rudiments of adventitious roots arise.
Regeneration. Lost parts of the body of Fungi and Algae are
152 BOTANY

often replaced, a direct regeneration thus taking place. In more

highly organised plants such a proceeding is extremely rare. It
occurs most readily in embryonic organs, such as growing points, when
portions have been lost, and is most often found in seedlings. Thus
in seedling plants of Cyclamen even a severed leaf-blade has been
found to be replaced. As a rule, however, when regeneration
processes are requisite in higher plants and the necessary preformed
organs are not present in a resting or latent condition, the older
tissues return to the embryonic condition and give rise to new
143
growing points of shoots ( ). That this is a provision for the
indirect replacement of lost parts is confirmed by the fact that direct
regeneration is far more frequent among animals than plants.

The Phylogeny of the Internal Structure

The phylogeneticdifferentiation in the internal structure of a

plant does not altogether coincide with the progress of its external

Fir,. 160. Surface view and transverse section of the thallus of Marchantiapolymorpha. In A, an
air-pore, as seen from above ; in B, as seen in cross-section, (x 240.)

segmentation. Even unicellular plants of the Siphoneae may

exhibit a high degree of external differentiation thus the uni- ;

cellular Alga, Caulerpa (Fig. 295), has developed appendages

having outwardly the forms of leaf, stem, and root. Similarly, the
Red Seaweed, Hydrolapaflium (Fig. 9), although composed almost
entirely of one form of cells, bears in its external segmentation a
resemblance to one of the most highly organised plants. The
internal differentiation of this Alga has only advanced so far, that
the outer cells containing the red chloroplasts form an assimilat-
ing tissue of isodiametrical cells, while the internal colourless and
SECT. I MORPHOLOGY 153

more elongated cells function as a conducting tissue. The relatively

highest degree of internal development found in the Algae is attained
by the Laminariae. In their stem-like axis, which may have a consider-
able thickness, the external tissues frequently contain canals filled
with mucilaginous matter ; while internally are found rows of cells
108
resembling sieve-tubes ( ). The axes themselves grow in thickness
through the continuous division of the cells of one or more of the
outer cell layers. A kind of cortical tissue is formed as a result
of this growth which exhibits concentric layers the innermost ;

cells gradually elongate

and pass over into the
so-called medulla. In the
larger Fungi internal differ-
entiation is the result of
the more or less intimate
union of the intertwining
hyphse. In extreme cases
the hyphae forming the f \

body of the Fungus may

be closely woven to-
so
gether as to give, in a
cross-section, the impres-
sion of a parenchymatous
tissue (Figs. 106, 107), in
which, on the subsequent
thickening of the cell walls,
the pits inadjoininghyphae
coincide. In the fructifica-
tions of many of the Hy-
menomycetes and Gastero- Flu. 161. Transverse section of tlie stem of Mnium undu-
mycetes, someof thelonger latum. I, Conducting-bundle ; c, cortex ; e, peripheral
and more swollen hyphae cell layer of cortex ; /, part of leaf ; r, rhizoids. ( X 90.)

contain a homogeneous or
cloudy, highly refractive, and, in some instances, coloured substance,
and appear, accordingly, to serve as a special tissue for the purpose of
conduction. A marked advance in the differentiation into different
tissue systems is first apparent in the Bryophytes, but even in them
the formation of an epidermis distinct from the fundamental tissue is
exceptional. In the thallus of the Marchantieae and on the spore
capsules of the Anthocerotaceae and Bryineae, the external layer of
cells become more or less sharply defined from the underlying tissues.
In the Marchantieae (Fig. 160) this outer layer is pierced by openings
which have been termed breathing-pores, but these have a different
origin from the stomata of higher plants. They are rather, as
144
LEITGEB ( )
has shown, openings into cavities, which have arisen
through the overarching of certain portions of the surface by other
154 BOTANY PA3CI i

more rapidly growing portions. In the Anthocerotaceae and Bryineae,

on the other hand, stomata similar in structure to those of the
Pteridophytes and Phanerogams are found in the outer cell layer of
the spore capsules. It would seem, however, that these stomata of
the Bryineae are probably not homologous with those of higher
plants. It is more reasonable to regard them as merely analogous
formations, such as from internal causes so often occur in the
evolution of organs. In the stems of many of the Bryineae there is
also developed a simple form of conducting bundle (Fig. 161); and
the many-layered midrib of the single -layered leaf lamina is al>i>
traversed by a conducting strand. In spite of their more advanced
differentiation, the Bryophytes may still be included, just as they
were originally in 1813 by AUGUSTE PYUAME DE CANDOLLE (
14
"'),
in
his classification of the vegetable
kingdom according to the natural
system, with the other lower Cryptogams in the class of CELLULAR
PLANTS, as distinguished from the VASCULAR PLANTS or Pteridophytes
and Phanerogams. A separation of the tissues into the three
systems of epidermal, fundamental, and vascular tissue occurs for
the first time in the vascular plants associated with the development
of roots ;
while the systems themselves also exhibit a widely extended
division of labour.

The Ontogeny of the Internal Structure

Just as the phylogenetic development of the vegetable kingdom
has proceeded from the simple to the complex, every plant in its
ontogeny passes through a corresponding development. The study of
the ontogenetic development is termed embryology. The early stage
of a new plant body is called the embryo or germ, and its further
development spoken of as germination.
is

However a plant may arise, whether from an asexually produced

spore or from a fertilised egg, its first inception is always as a single
cell. In unicellular, spherical, or rod -shaped organisms, such as
Gloeocapsa polydermatica (Fig. 1) or Bacteria (Fig. 4), the whole course
of development is concluded with the cell division which gives birth
to two new independent organisms (Fig. 1). If the cell divisions be
continuous and parallel, and the newly developed cells remain in
contact, CELL FILAMENTS (Fig. 4 a*) will be formed if the division
;

walls have different inclinations, and are at the same time all in the
same plane, CELL SURFACES are produced and if the walls are formed
;

in three dimensions of space, CELL MASSES are the result. Such an

organism will attain but a low degree of development if all its cells
have a like value. With the distinction into BASE and APEX a plant
manifests a higher degree of differentiation. A VEGETATIVE or
GROWING POINT is then developed, usually at the apex (Fig. 5).
This retains its embryonic nature, while the parts produced from it
 MORPHOLOGY 155

attain a mature condition, a distinguishing feature of the ontogeny

of plants as compared with animals (p. 4). In more simply
constructed plants the grow-
ing point consists of a single
cell. This assumes more
and more the character of
an APICAL CELL from which
all the organs of the plant
take their origin ; thus, in
the case of Cladostephus ver-
ticillatus (Fig. 7), the many-
celled main axis terminates
in a single conical cell which,

by transverse and longi-

tudinal divisions, gives rise
to the cellular system of
the whole plant. Its lateral
branches are likewise formed
from similar apical cells,
which develop, in regular
acropetal order, from certain
of the lateral cells of the
parent stem, and determine
the character of the branch-
The growing point of Di<:tyotH dichotoma, show-
ing. Flat, ribbon-like l-'ic;.li'c'.

ing the dichotomous branching. A, Initial cell. (After

plants also, such as Dicty- E. DE WlLDEMAN, X 500.)
ota dichotoma ( U6 ) (Fig. 8),

may have conical but correspondingly compressed apical cells (Fig.

162 A], from which segments are cut off by concave cross walls,
and become further divided by subsequent
longitudinal walls. The dichotomous
branching so apparent in Dictyota is pre-
ceded by a longitudinal division of the
apical cell into two equal adjoining cells
(B, By
a, a).the enlargement and con-
tinuous division of these two new apical
cells the now bifurcated stem becomes
prolonged into two forked branches. In
represent .,.
other ribbon-like Algae, on the other hand,
tion of the apex of Metzgena fur- and in similarly shaped Hepaticae, as in
f i a an d Awum (v
cato in process of branching, viewe.1 147
Metzger ),
the apical cell
from the tlorsal side, a, Apical . . , ('
cell of parent shoot ; 6, apical cell IS wedge-shaped (V Ig. 1 63), and the SUCC6S-
of daughter shoot. (After KNY, s i ve segments are cut off
alternately right
and left by intersecting oblique walls ;

from these segments the whole body of the plant is derived by further
division. The apparently strictly dichotomous branching of Hepaticae
156 BOTANY PART I

provided with such apical cells is in reality due to the early develop-
ment of new
apical cells in young segments (Fig. 163 b). In the
case of the erect radially symmetrical stems of the Musci, most Ferns
and Equisetaceae, the apical cell has generally the shape of an inverted
three-sided pyramid ( 148 ) with a convex base, and forms the apex of
the vegetative cone characteristic of the more highly organised
plants. In the common Horse-tail (Equisetum arvense), for example,
the apical cell of the main axis viewed from above (Fig. 165 A),
appears as an equilateral triangle, in which new walls are successively

Kiii. 104. Median longitudinal section of the vegetative cone of Kyuisetvm arvense.
Explanation in the text. (X 240.)

formed in a spiral direction, parallel to the original walls (p). Each

new segment thus derived divided by a new division wall (Figs.
is

164, 165 m) into an upper and lower half; each of these halves, as
is shown most
clearly by an optical section just below the apical cell
(Fig. 165 B), becomes again divided by a sextant wall (s) into two
new cells. It is unnecessary to trace the further divisions, and it
will suffice to call attention to the fact, that all cell walls parallel to
the outer surface of such vegetative cones or portions of plants are
termed PERICLINAL WALLS, while such as meet the surface and the
periclinal walls at right angles are designated ANTICLINAL; of the latter
those in the plane of the axis of an organ are called RADIAL. Some
distance below the apical cell of Equisetum arvense the first leaf-whorl
arises from the vegetative cone as a circular ridge, which grows by
 MORPHOLOGY 157

the formation of cell walls inclined alternately inwards and outwards

in the wedge-shaped marginal cells (Fig. 164/). This is succeeded
at a lower level by other and older leaf-whoris (/', /"). An initial
cell (g) may be distinguished in the axil of the second leaf-whorl,
and this is destined to become the three-sided apical cell of a side
branch.
In the Lycopodinae, the most highly developed of thePteridophytes,
a distinct apical cell can no longer be recognised, while in the
Phanerogams the cells of the vegetative cone are arranged as shown
in the accompanying figure of Hippuris vulgaris (Fig. 166), in which

<l

Pio. 1C5. A, Apical view of the

vegetative cone of Equisetum
orven.se ; B, optical section Fio. 160. Median longitudinal section of th
of the same, just below the vegetative cone of Hippuris vulgaris. d, der-
apical cell ;I, lateral walls matogen ; pr, periblem ; pi, plerome ; /, leaf
of the segments. ( x 240. )
rudiment, (x 240.)

the embryonic tissues are arranged in layers which, as was first

noticed by SACHS ( ),
form 149
confocal parabolas. The outermost
layer, which covers both the vegetative cone and also the developing
15
leaves, is distinguished as the DERMATOGEN ( ) (d) ;
the cells of the
innermost cone of which the central cylinder terminates,
tissue, in
constitute the PLEROME (pi) while the layers of cells lying between
;

the dermatogen and plerome are called the PERIBLEM (j>r). In the
same figure may be noticed the uniformity with which the dividing
walls of the different layers intersect at right angles. This arrange-
ment was regarded by SACHS as characteristic of the whole plant
structure. The anticlinal walls at right angles to the surface form a
system of orthogonal trajectories with the periclinal walls.
While SACHS regarded the insertion of new cell walls at right angles to pre-
existing ones as the principle of their succession, BERTHOLD and EIIREEA ( 151 ) have
endeavoured to show that the curvature and mode of insertion of a partition wall
168 BOTANY

obey the same laws as do weightless films of liquid. The cell wall tends at the
moment of its origin to assume the same position as a film ia a mass of soap-suds
would under similar conditions.

True ROOTS are first found in the Pteridophytes, and in them

1;V
possess an apical cell in the shape of a three-sided pyramid ( -) (Fig.
167 /). In addition to the segments given off by the apical cell
parallel to its sides, it also gives rise to other segments (k)

Ki<;. I'i7. Median longitudinal section of the apex of a root of 1'terix eretica.
t, Apicalcell : initial cell of root-cap
7>-, t, root-cap, (x -J40.)
;

parallel to its base. It is from the further division of these latter

cap-like segments that the ROOT-CAP is derived. In those ferns in

which the root may form a terminal shoot (p. 47) the apical cell of
the root according to the investigations of ROSTOWZEW ( 10;? ) forms the
apical cell of the new. shoot. It ceases to give off segments parallel
to the base, and the segments divide differently from those forming
the root. In the roots, as in the stems of the Lycopodinae, no apical
cells are found. In like manner the roots of Phanerogams, although
exhibiting several different types of apical growth, follow the same
law in the arrangement of their elements as the vegetative cones of
 MORPHOLOGY 159

the stems. It will, accordingly, be sufficient to describe a root of

one of the Gramineae (Fig. 168) as a representative of one of these
154
types ( ). The vegetative cone of this root differs from that of the
stem previously described (Fig. 166) in the possession of a root-cap.

Kio. Ki8. Median longitudinal section of the apex of a root of the Barley, Hordeum vulgare. If,
calyptrogen ; rf, dennatogen; c, its thickened wall pr, periblem pi, plerome en, endoUerniis
; ; ; ;

', intercellular air-space in process of formation;

cell row destined to form a vessel
, ; r,
exfoliated cells of the root-cap, (x 180.)

The dermatogen (d) and periblem (pr) unite at the apex in a single
cell layer,outside of which lies the CALYPTROGEX (k) or layer of cells
from which the root-cap takes its origin. In many other roots,
however, the formation of the root-cap results from the periclinal
division of the dermatogen itself, which, in that case, remains distinct
from the periblem. In the apices of Gymnosperms the dermatogen,
160 BOTANY PART i

periblem, and calyptrogen are not marked out as distinct regions.

In roots, as in stems, the plerome cylinder (pi) almost always
terminates in special initial cells.
At a short distance below the growing point the embryonic tissue
loses its meristematic character, and becomes transformed into the
differentiated body of the plant. As a general rule, in plants with
an epidermis, primary cortex, and central cylinder, the epidermis
is developed from the dermatogen ; the primary cortex from the
periblem the central cylinder from the plerome.
;
This differentiation
of the tissues does not take place in all cases; an already determined
separation of the several tissues must not be assumed to extend to
the embryonic tissue, the peculiar cell arrangement of which is due
rather to physical causes. The vascular bundles must pass through
the periblem in order to reach the leaves. The periblem is therefore
capable of producing, not only the primary cortex, but also the
vascular bundles and accompanying tissues of the central cylinder.
The terms dermatogen, periblem, and plerome are employed merely
for convenience to designate certain cell layers, and are not to be
regarded as significant of any peculiar histogenetic or tissue-forming
ability. The external layer from which the epidermis develops usually
remains a single cell layer. The rudiments of the still undeveloped
vascular bundles soon appear in the central cylinder as procambium
strands ; while the endodermis of roots is derived at an early stage
from the innermost layer of the cortex.
In stems with apical cells the RUDIMENTS OF NEW LEAVES and
SHOOTS are developed from single peripheral cells, or cell groups of
the vegetative cone (Fig. 164). In such cases, not only the new
shoots, but even the leaves, usually begin their development with an
apical cell. The apical cells of the leaves, however, soon disappear,
and further growth proceeds along their whole margin.

In those fern leaves which are able to form terminal buds (p. 31) GOKBEL ( 155 )
has observed the direct transition of the two-sided apical cell of the leaf tip to the
three-sided pyramidal apical cell of the young shoot ;
this takes place by the

origin of a suitably placed wall in the apical cell of the leaf.

In a stem with no apical cell (Figs. 17, 166), the rudiments of

the leaves and new shoots first appear as small protuberances, the
formation of which is generally initiated by the periclinal division of
a group of periblem cells while the cells of the overlying dermatogen
;

continue to divide by walls at right angles to the surface. In the

case of new shoots developing at some distance from the growing
point of the parent stem, the cells from which they are destined
to arise retain for that purpose their original embryonic character.
The formation of the buds on the rapidly growing shoots of bushes
and trees in the spring may be postponed, so that the rudimentary
lateral shoots first appear in the axils of the eighth or even the tenth
 MORPHOLOGY 161

youngest leaves, and consequently at points where the differentiation

of the surrounding tissue has
156
already begun ( ).
The vegetative cone, in the
case of strictly
dichotomously
branching shoots (cf. p. 19), in-
creases the number of its cells
in the direction of the
plane of
the subsequent bifurcation, and
eventually gives rise to two new
growing points.
With the exception of those
Pteridophytes whose roots as

FIG. 169. Thuja, nccidentalis. A, Median longitudinal

section through the ripe seed (x .5). B, C (x 2) ;

D, E (nat. size), different stages of germina-

tion ; h, hypocotyl ; c, cotyledons ; r, radicle ;

v, vegetative cone of stem.

well stems are dichotomously branched

as
(cf. p. the branches of all other normal
46),
roots, whether growing by means of an apical
cell or not, arise in acropetal succession ;
the
branching first begins in regions considerably
removed from the growing point, where the
differentiation of the tissues is already com-
plete. In Phanerogams new roots are developed
in the pericycle in Pteridophytes in the inner-
:

most cortical layer. The lateral roots must

consequently push through the whole cortical
layer of the parent root. They are situated
either directly opposite to the vascular strands
of the parent root, or between the xylem and

phloem strands. The number of rows of lateral

therefore, as VAN TIEGHEM (
157
roots is, )

pointed out, either equal to or double the

number of vascular strands. As the strands
of the vascular bundles of roots take a straight
Fio. 170. Seedling of CVi^.//. tu
Jietulits. h, Hypocotyl ; c,
course, the lateral roots must similarly form
cotyledons ; hu; main root ;
straight rows. The distances between the rows
sw, lateral roots root-
; r,
themselves are either equal, or, when the lateral
hairs; c, epicotyl; I, I', foliage
leaves. (Nat. si/e.) roots are situated to the right and left of each
vascular strand, the rows are arranged in
pairs with wider intervals between each pair.
M
162 BOTANY

The stage
of the ontogeny of the plant known as GERMINATION
is specially defined in Phanerogams, where the term is limited to
the further growth of the embryo already present within the
seed. The embryo, enclosed within the seed, by the time the latter
is shed from the parent plant generally exhibits the segmentation
characteristic of Cormophytes. Protected by the hard seed-coats, it
is enabled to sustain a long period of rest. Abundant deposits of
nutritive material in the
embryo itself, or surround-
ing it, are provided for its
nourishment during ger-
mination. The different
segments of a phanerogamic
embryo have received dis-
names thus, as in
tinctive ;

theembryo of the American

Arbor Vitae (Thuja occiden-
talis, Fig. 169), the stem
portion (h) is termed the
HYPOCOTYL, the first leaves
(c) are the SEED LEAVES
or COTYLEDONS, while the
root (r) is distinguished as
the RADICLE. The tap-root
of the fully-developed plant
is formed by the prolonga-
tion of the radicleand later
gives rise to lateral roots.
FIG. 171. Seedling of Acacia jiycnantha. The cotyledons J n ~p j 170 a **
germinating
have been thrown off. The foliage leaves 1-4 are . P" . _ ,

pinnate, the following leaves bipinnate. The petioles plantlet Ol the Hornbeam
of leaves 5 and 6 are vertically expanded ; and in the
(CdTpinUS jBetulus) is shown
following leaves 9, modified a.s phyllodia, bearing

nectaries, n.
^8
(About J natural size.)
j fch j
hypocotyl
J r
fc
J /)/ an(J
\

cotyledons (c)two its ;

radicle has already developed into a tap-root (hw) with a number of

lateral roots (sw). An internode and foliage leaf (I) have been
produced from the vegetative cone of the stem while the next higher ;

internode is also distinguishable, but has not yet elongated, and a

second foliage leaf (I 1 ) is unfolding.
A highly organised plant, which begins its development with the
simplest stages and gradually advances to a higher state of differentia-
tion, to a certain extent repeats in its ontogeny its phylogenetic
development. In the process of its ontogenetic development much
has been altered, and much omitted, so that it presents but an
imperfect picture of its past history ; nevertheless, this representation
is valuable, and, next to comparative methods, furnishes the most

important source of our morphological knowledge. Whatever is true

 MORPHOLOGY 163

of the development of a plant from the embryo is also, as a rule,

applicable to its further growth from the growing point, and, conse-
quently, a knowledge of the mode of development at the growing
point is of great importance in establishing homologies. The earlier
a characteristic makes itself apparent in the embryo, or the nearer it
is to the growing point of the old plant, so much the greater is its

value in determining the general relationships existing between the

different plants ; the later it is exhibited in the embryo, or the
farther removed it is from the growing point of the plant, the less
its general value, but the greater, in proportion, its importance in

defining the character of a genus or species. From the fossil remains

of former geological periods, it is safe to conclude that such Conifers
as Thuja, Biota, and the various Junipers, that now have scale-like
compressed leaves, have been derived from Conifers with needle-
shaped leaves. This conclusion is confirmed by the fact, that on the
young plants of the scaly-leaved Conifers typical needle-shaped leaves
are at first developed. The modified leaf forms do not make their
appearance until the so-called JUVENILE FORM has attained a certain
age, while in some Junipers needle-shaped leaves are retained through-
out their whole existence ( 158). Even still more instructive are the
Australian Acacias, whose leaf-stalks become modified, as phyllodia
(p. 45), to perform the functions of the reduced leaf-blades. The
demonstration of such an assertion is furnished by a germinating
plant of Acacia pycnantha (Fig. 171), in which the first leaves are
simply pinnate, and the succeeding leaves bipinnate. In the next
leaves, although still compound, the leaf-blades are noticeably reduced,
while the leaf-stalks have become somewhat expanded in a vertical
direction. At length, leaves are produced which possess only broad,
flattened leaf-stalks. As many other species of this genus are pro-
vided only with bipinnate leaves, it is permissible on such phylogenetic
grounds to conclude that the Australian Acacias have lost their leaf-
blades in comparatively recent times, and have, in their stead,
developed the much more resistant phyllodes as being better adapted
to withstand the Australian climate. The appearance, accordingly,
of the phyllodes at so late a stage in the ontogenetic development of
these Acacias is in conformity with their recent origin. Lathyrus
aphaca 44) the leaves of which in the mature plant are transformed
(p.
into tendrils, has the first leaves of the seedling provided with
leaflets. It may, in like manner, be shown that in the case of

plants with similarly modified leaf forms, the metamorphosis of the

leaves does not take place until after the cotyledons and the first
foliage leaves have been developed, and it is then usually effected
by degrees.
In the lower divisions of the vegetable kingdom, especially among
the Fungi, PLEOMORPHISM is a phenomenon of frequent occurrence.
In the developmental cycle of the species more than one living being
164 BOTANY PART I

or biont is found, and the bionts are characterised by differences in

form. Thus in the Rust of "\\r heat (Pucdnia graminis) the individuals
which live on the Barberry produce as reproductive organs aecidia
and spermogonia. From them arise the quite distinctly organised
biont living on the leaves and stems of grasses which is reproduced
by means of uredospores and teleutospores. From the latter, on damp
soil inthe spring, tubular germ-plants arise which in their turn bear
a speial kind of reproductive cell, the basidiospores. With this type
of spore the developmental cycle of the species is completed, since
from the basidiospores are again formed the individuals which live
on the Barberry.

In treating of the cell processes which constitute fertilisation (p. 92), it has
already been seen that they result in a doubling of the number of chromosomes
present in the nucleus. Doubtless, the process of fertilisation has repeatedly
originated in both animal and vegetable kingdoms as soon as a certain grade of
organisation had been attained in the course of phylogenetic development. In
every case the process of fertilisation must have resulted in a doubling of the
number of chromosomes. The single chromosome number must be assumed to be
the number originally occurring in the organism. According to the view of most
investigators the chromosomes are persisting entities in the nucleus. Further,
there is nmch in favour of the view that they are different from one another.
Under these conditions the doubling of the number of chromosomes in the act of
fertilisationwould lead to a progressive increase in their number if the reduction
division (p. 84) had not been introduced into the course of development of the
organism in this division the number is reduced to one half, that is to say, a return
;

is made to the original condition. If in the ontogeny of an organism, the reduction

division follows immediately on fertilisation all the bionts of the species will show
the single number of chromosomes. This was, in any case at first after the distinc-
tion of sex, always the condition of things, and it is found in many existing
Thallophytes. In the further course of phylogenetic development the sexual
product developed into a special biont, which accordingly had the double number
of chromosomes in its tissues. Not until the formation of its reproductive cells did
this biont proceed to the reduction division by which the number of chromosomes
returned to the original single number. When in the developmental cycle of a
species bionts with the single chromosome number and others with the double
number are represented, we have to do with an alternation of generations. Such
an alternation of generations is found in all Bryophyta, Pteridophyta and Phanero-
gams, i.e. in plants from the Bryophytes onwards. The generation with the
double number of chromosomes became in these plants the dominant one, while
that with the single number underwent a progressive diminution in size. Thus
the plant body which is most prominent in Pteridophytes and Phanerogams is the
generation with the double number of chromosomes, while the corresponding
generation with the single number becomes so inconspicuous that it ultimately
requires to be carefully looked for. This is especially the case for the Phanero-
gams, in which this generation no longer has an independent existence, but
completes its development within the generation with the double number of
chromosomes. The generation with the single number of chromosomes is in
plants termed the gametophyte ;
that with the double number the sporophyte.
In order to extend the distinction to the animal kingdom the terminology
 MORPHOLOGY 165

must be altered the organism with the single number of chromosomes may be
;

termed the haploid, or haploid generation, that with the double number the
diploid, or diploid generation.
The union of paternal and maternal chromosomes in each sexual act results in
the nuclei of the offspring being provided with maternal and paternal chromosomes
in equal numbers. All the nuclei of a diploid generation contain each kind of
chromosome in duplicate in two equivalent chromosomes. In the reduction division
by which the formation of the reproductive cells is prepared for, the chromosomes
become united (as hag been seen on p. 84) in pairs during the prophase.of the.
division. It is to be assumed that the pairs are formed of the corresponding

parental chromosomes. In the separation which follows during the metaphase of

the reduction division each chromosome may apparently pass to either of the
two daughter nuclei, but always in such a manner that the chromosomes of -each
pair pass to different nuclei. Thus on the one hand, all combinations of the
parental chromosomes are possible in the products of this division on the other
;

hand a chromosome of eacli kind is ensured to each of the resulting nuclei. All
these morphological processes, which are matters of observation, lend great support
to the assumption that the nuclei are the bearers of the inherited properties of the
159
organism ( ).

By those apogamous processes which exclude a reduction division (p. 93) an

alternation of generations is also abolished.

16
Structural Deviations ( )

Plants, even of the same

species, never exactly resemble each
other. Every individual organism has its own peculiar characteristics
by which it may be distinguished from every other of the same
species. To a certain extent these characters may be due to
ATAVISM, or the reappearance of previous ancestral qualities. Most
individual deviations belong, however, to the so-called FLUCTUATING
VARIATIONS. These occur in all species, and may be compared to the
excursions of apendulum to either side of its position of equilibrium.
Abrupt and striking deviations of the individual from the type of
the species are called MONSTROSITIES. When these induce a dis-
turbance of function a diseased condition results. The causes of
monstrositiesmay lie in the organism itself ; in other cases they can
be shown to stand in relation to external causes. Among EXTERNAL
CAUSES OF VARIATIONS the influence exercised by parasites upon the
development of the portions of the plant is
particularly striking.
Euphorbia Cyparissias, when attacked by a rust fungus (Aecidium
Euphorbiae), becomes sterile, remains unbranched, has shorter and
broader leaves, and in its whole appearance is so changed as scarcely
to be recognisable. Plant lice sometimes cause a flower to turn
green, so that instead of floral leaves green foliage-like leaves appear.
Another peculiar example of abnormal growths is afforded by the
GALLS or CECIDIA produced on plants by Fungi, or more frequently by
insects, worms, and arthropods. The effect of these formations on the
166 BOTANY PART I

normal development of the tissues of a plant is more or less dis-

turbing, according to their position, whether it be in the embryonic
substance of the growing point, in the tissues still in course of
differentiation, or finally in those already developed. The larvae of
Ceddomyia rosaria live in the growing points of Willow stems, and
occasion a malformation of the whole shoot by the production of
"
galls, known as willow-roses," which are composed of modified
leaves and axes. Flies (Diptera) often deposit their eggs in the
tissues of partially developed leaves, in consequence of which the
leaves become, according to their age when attacked, more or less
swollen and twisted. After the leaves of the Oak have attained
growth they are often stung by a gall-wasp of the genus
their full
Cynips. The poison introduced by the
sting, and also by the larvae
hatched from the eggs deposited at the same time, occasions at first
only a local swelling of the leaf tissue, which, finally, however,
results in the formation of round, yellow, or red galls on the lateral
ribs on the under side of the leaf. As galls differ materially from
one another according to the nature and cause of their formation, it
is
generally possible to determine the particular exciting cause to
which they are due.
In the monstrosities resulting from internal causes it is also the
case that the earlier they commence the more extreme they tend to
be. When the embryonic substance of the growing point is affected
by such an influence altogether unexpected modifications of the
usual order of growth may result. In other cases intermediate
structures of more or less monstrous form result. As the embryonic
substance of the growing point is of itself capable of producing all
such forms as are peculiar to the species, instead of a flower a
vegetative shoot may be developed, or the growing point of a root
may continue its further development as a stem. Leaves, even when
somewhat advanced in growth, may under changed conditions vary
their usual character, particularly within the limits of their possible
metamorphosis ; for example, the staminal and carpellary leaves of a
flower may thus become transformed into additional perianth leaves.
The later the rudiments of an organ are acted upon by a disturbing
influence, so much the less far-reaching are the modifications which
are produced ; thus intermediate forms between two organs may
be produced which correspond more or less closely to one or the
other of them. Neither the abnormal interchangeability, at times
manifested between morphologically different members, nor the
development of intermediate forms between them, can be considered
as proof of their phylogenetic connection. MALFORMATIONS ARE,
ACCORDINGLY, TO BE ACCEPTED AS EVIDENCE IN MORPHOLOGICAL
QUESTIONS ONLY IN THE RAREST CASES. When, as is only exception-
ally the case, they represent reversions, this value may be attached
to them.
 MORPHOLOGY 167

The study of the abnormal formations and functions of the plant is called
PHYTOPATHOLOGY. When the pathological structures are alone in question it may
he termed pathological morphology. Malformations which affect the external form
are treated by themselves as Vegetable Teratology (Phytoteratology).
still Patho-
logicalchanges in the form and contents of the vegetable cells and tissues form the
subject matter of pathological plant anatomy.
The study of the aetiology of abnormal vegetable tissues shows that various
external influences may give rise to them the most important causes are the in-
;

fluence of wounding and of the attacks of parasites. According to EUNST KUSTKU

abnormal cells may be characterised by their contents, the thickening of their walls,
or their unusual size (hypertrophy) ; in the last case they may also show
by
divisions which would not normally occur. The abnormal tissue may resemble
that from which it originates or deviate more or less widely from it. The tissues
formed in the healing of wounds (cf. p. 151) and those of galls may be mentioned
as examples. The tissues of galls are characterised more by the increase in
number than by the size of their cells, which may undergo the most various
modifications. There is often a considerable accumulation of albuminous substances
and carbohydrates in these tissues. A modified part of a plant in many cases
appears as if due to an arrested development. Dwarf forms and defective
differentiation of tissues with its accompanying impairment of function are

examples of this. In all cases in which the cells of a tissue no longer present their
normal properties they are liable to undergo sooner or later regressive changes
leading to degeneration and necrosis.
 PART I

GENERAL BOTANY

SECTION II

PHYSIOLOGY
 SECTION II

PHYSIOLOGY

PLANTS, like animals, are living organisms. Beginning their develop-

ment with the simplest structure, and increasing in size from internal
causes, they assume their definite form and complete their existence
according to laws determined by inheritance. Surrounded by a
world, which differs very widely from them as regards chemical
constitution, they produce the substances necessary to their growth
from the raw materials afforded by the environment. To this end
the different parts of their bodies are enabled by independent
movements to take such relative positions as are most favourable to
their mode of growth. Like other living beings plants sooner or
later perish. In spite, however, of the limited duration of life and
number of individuals, the continuance and extension of the species
by an ability to reproduce like organisms.
are provided for
NOURISHMENT, INDEPENDENT GROWTH, POWER OF MOVEMENT, and
REPRODUCTION are, together with RESPIRATION, the most striking
attributes which characterise plants as living organisms, and distinguish
them from all lifeless bodies.
An organism consisting of but one cell, as is shown by the life of
the simplest plants, is capable of exercising all the functions necessary
for the continuance of its existence. In the case of plants, however,
which consist of many hundreds or thousands of cells arranged in
three dimensions of space, it is impossible, for purely physical
reasons, that all the cells should bear the same relations to the outer
world. The cells in the interior must exist under conditions
altogether different from those which are in direct contact and inter-
course with the world outside. Consequently, the differently arranged
elements must be adapted for different modes of life, and, since they
exercise their functions in different ways, they show what is called
DIFFERENTIATION.
This necessary division of labour has led to the development of
external organs and internal structures wonderfully adapted to the
requirements of the whole plant. Correlated with the natural
groups of plants, there are certain differences as regards form
and function. In all plants, however, those organs to which the
171
172 BOTANY PART i

same functions are assigned have assumed the form most efficient
for their purpose so that, for example, the leaves and roots of plants
;

otherwise most dissimilar are constructed on the same general plan.

In proof of this may be cited the general terms leaf, root, stem, and
flower, the comprehensiveness of which is even more evident in
popular speech than in the technical language of Botany, which has
given to these terms a more strictly defined and limited meaning.
Similarity in the appearance and structure of organs indicates the
exercise of common functions and duties ; while dissimilarities in the
form and structure of different organs such as the leaf and root are
indicative, .on the other hand, of their different utility to the plant.
There lies, then, in the morphological and anatomical development of
an organ an unmistakable proof that it exists because of its function,
and that it is not of accidental origin.
The attributes and functions of organs, as well as of single cells,
are the subjects of physiological study.
It is evident, however, that
such study must be based upon an intimate knowledge of the outer
and inner structure of plants ; just as the working and efficiency of
a machine first become comprehensible through a knowledge of its
construction. The principal and most reliable source of physiological
knowledge however, obtained by experiment.
is,
It is the province of
Physiology to discover the points of corre-
spondence among the numerous individual phenomena, and to bring
to light such as possess an essential functional significance. On the
other hand, it is the variations, or family peculiarities, which are of
value in Systematic Botany, since from them a knowledge of family
relationships may be derived. For example, it suffices for the
physiological understanding of flowers to know that they are the
organs of sexual production in higher plants ; that the male cells are
somehow developed from the pollen formed in the anthers ; that from
the female cell enclosed within the ovules, after its union with a
male cell from the pollen-tube the embryo or rudimentary plant is
derived. These important facts are equally true for all flowers, no
matter how dissimilar they may appear.
Physiology considers peculiarities of form in so far as they are of
service for special purposes arising from the relation of the plant to
its
surroundings (Pollination, Distribution of Seeds, Water-plants,
etc.). This side of physiology is termed BIOLOGY, or better (ECOLOGY
(Bionomics) since the former term is often applied to the science of
living beings in general.

The Physical and General Conditions of Life and Vital

Properties of Plants

With the exception of the more or less fluid developmental stages

in some of the lower organisms, as in Amoeba or the plasmodia of
SECT, ii PHYSIOLOGY 173

Myxomycetes (p. 54), plants, in spite of the great amount of water

contained in them, are of the nature of solid bodies. As such they
possess in common with inanimate objects the physical attributes of
weight, rigidity, elasticity, conductivity for light, heat, electricity, etc.
Important as these attributes are for the existence and continuance
of the life of a plant, they do not constitute that life itself.
VITAL PHENOMENA ARE ESSENTIALLY BOUND UP WITH THE LIVING
PROTOPLASM. No other substance exhibits a similar series of remark-
able and varied phenomena, such as we may compare with the
attributes of life. As both physics and chemistry have been restricted
to the investigation of lifeless bodies, any attempt to explain vital
phenomena by
solely chemical and physical laws could only 1>e induced
by a false conception of their real significance, and must lead to fruit-
less results. The physical attributes of air, water, and of the glasses
and metals made use of in physical apparatus, can never explain
qualities like nutrition, respiration, growth, irritability and repro-
duction. It would, indeed, be superfluous to emphasise the fact, were
it not that this error is from time to time repeated.

The phenomena of life can only be studied and determined by the

most careful observation and critical examination of living organisms.
It is therefore necessary to establish what part the purely physical
and chemical properties which belong to all bodies, take in the
phenomena of life, and to what extent they are essential to the
maintenance of life itself. A perception of the strictly physical and
chemical processes going on within an organism is especially desirable,
though their exact determination is not more important than that of
the vital reactions, because operations are then involved with the
causes and effects of which we are already familiar. In questions
regarding strictly vital phenomena the case is quite different ; for it

then becomes impossible to predict what effect a particular cause will

produce.

The free end of a horizontally extended flexible rod bends downwards merely
by itsown weight. The same result will follow if any part of a dead plant, such
as a dry stem, be substituted for the rod. But if a living, growing stem be used
in the experiment, then the action of gravity will manifest itself in a manner
altogether at variance with its ordinary operation. That part of the stem which is
still in a state of growth will ultimately curve upwards, and BY ITS OWN ACTIVITY

ASSUME AN UPRIGHT POSITION it moves in a direction exactly contrary to the

;

attractive force of gravity. If a tap-root be similarly

experimented upon, it will,
on the contrary, continue its downward movement until it places itself in a line
with the direction of the attraction a rhizome, however, under like circumstances,
;

would constantly maintain its growing apex in a horizontal position. In these

three experiments the force of gravity is exerted upon horizontal portions of
plants. The physical conditions are the same in each case, yet how entirely
different the results!

The explanation of this dissimilarity in the effects of the action

174 BOTANY PAKT i

of gravity is to be sought in the fact that gravity acts upon living

substances, not only physically but also in another way, as a stimulus
which induces a response in the internal activities of the plant body.
In these particular experiments it is the processes and forces of growth
which are locally increased or hindered by the action of gravity, and
produce results which do not correspond either qualitatively or
quantitatively with the known operations of the laws of gravity.
Living substance is dominated by the operation of stimuli. Irritability
is its most important attribute, for it is irritability alone that renders

possible what we call life.

By irritability is meant the undoubted, though not fully under-

stood, causal between a particular influence and the
connection
peculiar vital response of an organism. The disproportion that may
exist between an influence and its ultimate effect is plainly apparent in
a steam engine or in the firing of firearms. The slight pressure of the
finger in firing a cannon has as little correspondence, either quantita-
tively or qualitatively, with the destructive effect of the shot, as the
small effort necessary to open the throttle- valve of a locomotive has to
the continuous motion of a heavily laden goods train. The opening of
the valve of an engine before the steam is up has no effect ; it is only
when, by this process, the compressed steam is liberated that it is
followed by such enormous results. In the engine the connection
between the external influence and its effect is known ; in the effects
of stimuli on protoplasm this connection is not apparent, for in the

protoplasm the intermediate processes remain invisible to the eye,

even when aided by the best microscope. There is, however, no
occasion for the supposition that the connection between the
stimulating cause and its effect on the protoplasm is accomplished
by processes which are otherwise foreign to the protoplasm itself, and
which are called into existence only under the influence of a special
force, the vital force. It was formerly thought necessary to ascribe
not only all indications of life, but even all the transforming processes
carried on within animate objects, to the effects of a special vital force.
Now, however, the conception of the vital processes has become so
modified as no longer to require the supposition of such a special
vital force ; while the impossibility of explaining the manifold variety
of their manifestation by the action of a single force, and the advances
made in chemistry (cf. pp. 4, 5), have shown the futility of such a
supposition.
Although, at the present time, the existence of a special, inde-
pendent vital force is denied by Physiology, and only such agencies
are accepted as are inherent in the substance of an organism itself,
still we must at the same time take account of such a vital force in

so far as it be regarded as the expression of a living substance,

may
endowed with a peculiar, internal structure, which is in some way so
constituted that certain actions and conditions are followed by definite
SECT, ii PHYSIOLOGY 175

vital processes. It is, then, this peculiar quality of irritability

that distinguishes living protoplasm from other bodies, and con-
stitutes the fundamental distinction between living and dead proto-

plasm. Such a view is, however, not contrary to accepted ideas ;

simple chemical bodies, indeed even chemical elements, such as

" "
sulphur, phosphorus, etc., exist in different modifications with
fundamentally different peculiarities. In considering living organ-
isms, it is the irritable or living modification of the protoplasmic
substance which must occupy the attention. The object, therefore,
of Physiology consists principally in discovering the attributes and
characteristics incident to the modifications of living protoplasm.
These attributes and characteristics are so distinctive as to
separate by a wide gap living bodies from all other matter. It is,
in fact, impossible to form any conception of the manner in which

living bodies have arisen on this once molten planet from lifeless
matter. Acceptance of the theory of evolution authorises, it is true,
the transfer of the inception of life on the earth to geological periods
separated by millions of years from the present time; but the character
and condition of origin of such dawning life remain no less incompre-
hensible. From a consideration, however, of the attributes of the
living substance, it can with safety be said that the external conditions
of life could not at that time have been so very different from those
now existing on the earth ; for it is a characteristic quality of living
matter that its vital activity, even its very existence, is circumscribed
and limited by external, cosmic influences. The
vitality of vegetable
protoplasm can only be preserved within a definite range of tempera-
ture, while its full vital activity is restricted to still narrower limits.
Too intense light or too little warmth destroys its life ;
while the
most minute quantities of certain poisons suffice to shatter instantly
and irrevocably that mysterious structure, in which, under favour-
able conditions, lies concealed the capacity to populate the whole
world.
Although living plants are themselves responsible for the manner
in which their vital phenomena manifest themselves, they stand,
nevertheless, in the closest reciprocal relations with their environment,
upon the condition of which they are dependent. From the outer
world they obtain not only their nourishment, but receive also from
it, particularly from the vibrations of light and heat, the energy that

they again expend in the manifold processes of their vital phenomena.

It is to the operation of these external influences that the stimuli
are due, which constantly call forth in vegetable protoplasm the
manifestation of vital phenomena. These external influences, how-
ever, are only serviceable to the processes of life when they operate
within definite limits of intensity. The lowest limit of intensity for
the effective operation of an external influence is designated the
MINIMUM, the highest the MAXIMUM, while that degree of intensity
176 BOTANY

at which it is operative in calling forth the most active manifestation

of a definite vital phenomenon is termed the OPTIMUM. Minimum,
optimum, and maximum are not, however, points of fixed value for
the organism, but vary, on the one hand, with the duration of
exposure on the other, with the sum of the other conditions.
;
For
the different vital processes of the same plant, and also for those
of distinct plants, these so-called CARDINAL POINTS are generally
different.

The cardinal points for the vegetation of cold climates are on the average much
lower than those of plants inhabiting temperate or tropical countries. The dis-
tribution of plants over the surface of the earth is thus to a considerable extent
dependent on their cardinal points. The highest are not, however, possessed by
tropical plants, but by small Algae and Bacteria which inhabit hot springs, the
water of whicli has a temperature of 70-80 C. The albumen of a hen's egg is
quickly coagulated by the water in which these plants find their suitable habitat,

Fio. 172. Protonenia of Schistostega osmundacea. The lens-shaped cells concentrate

the feeble light available in the habitat of the plant upon the chloroplasts.
(Cf. Fig. 214.)

while some thermogenic Bacteria can raise their own temperature to 70 C., and
even higher.
Some plants flourish best when exposed to bright sunlight, while the shade-
loving plants only attain their perfect development in a subdued light, such as
that of a forest, or like Schistostega in crevices and caves (Fig. 172). Not only
does the intensity of the required illumination differ for different species of plants,
and also for individuals of the same species, but the optimum intensity may change
as the development of the individual plant proceeds. Shade is absolutely essential
formany tropical plants in a young state, although at a later age they can endure
and may even require the full light
of the tropical sun. Young plants require to
be protected by specially planted shade -producing trees (Albizzia moluccana,
A. stipitata, etc.).
On exposure to a
low temperature, about the freezing-point of water, many
plants become frozen and generally die. Very sensitive plants may even become
frozen at a temperature considerably above zero, before ice has been formed in
their tissues. In the case of other plants the internal formation of ice in the
tissues does not of itself occasion death. The formation of ice always begins in
the intercellular spaces and not within the cells. Its continued formation is

accompanied by an increasing concentration of the cell sap ;

as a consequence of
 PHYSIOLOGY 177

this, ice firstbegins to form iu plants at a tenlperature below zero, and only
gradually increases in case of a greater reduction of temperature (according to MEZ,
to - 6). Such plants as inhabit Arctic and Alpine regions are able to recover
from the completely frozen condition. The power which the lower organisms have
of withstanding extremely low temperatures is very striking. In PICTET'S investi-
gations Diatomaceae were found to sustain a long exposure to a temperature of
-200 C. MACFADYEN and others have shown that many Bacteria can endure a
;

week's exposure to 250 of cold produced by means of liquid air and hydrogen ( J ).
A sudden change of temperature leading to the complete thawing of frozen

FIG. 173. Trees on the coast of Denmark ; to the left Prunus spinosa, to the right L'rut<ir<iu.i
oxycantha, the form of both being similarly modified by the influence of the sea wind. (After
WARMING : from SCHIMPER'S Plant-Geography.)

plants is better borne than repeated transitions from freezing to thaw. A long,
keen frost is therefore, as a rule, less injurious to vegetation than less severe
frost alternating with frequent partial thaws.

When the vital powers of reaction of the plant are opposed to

purely physical influences, it will naturally depend on their respective
strengths which shall become the dominating influence. A centrifugal
force a thousand times greater than that of gravity is powerless to
influence the reaction of the plant. The degree to which constant
and strong currents of air, such as sea-winds, may completely alter the
2
characteristic growth-forms of trees ( )
is illustrated by Fig. 173.
N
178 BOTANY

I. The Stability of the Plant Body

One most important and essential physical attributes of a

of the
plant is its Without that quality plants could retain no
rigidity.
enduring form. Plants which reach a considerable height and by
branching expand freely in the air are specially dependent on the
rigidity of their bodies so as to remain fixed in the position they
have attained. The capacity to return, by their own independent
movement, to favourable positions from which they may have been
forcibly displaced by external influences, is, in trees and shrubs, and
also in the more rigid herbs, restricted to the extreme tips of the
growing branches.
How great are the demands made upon the stability of plants will
be at once apparent from a consideration of a Rye haulm although ;

it is
composed of hundreds of thousands of small chambers or cells,
and has a height of 1500 mm., it is at its base scarcely 3 mm. in
diameter. The thin stems of reeds reach a height of 3000 mm. with
a base of only 15 mm. diameter. The height of the reed exceeds by
two hundred times, and thatof the rye haulm by five hundred times,
the diameter of the base. In addition, moreover, to the great
disproportion between the height and diameter of plants, they
often support a heavy weight at the summit ; the Rye straw
must sustain the burden of its ears of grain, the slender Palm the
heavy and wind-swayed leaves (which in Lodoicea seycJiellarum have a
length of 7 m. and a breadth of 3-4 m.), while at times the consider-
able weight of the bunches of fruit has also to be considered.
In plants, however, the rigid immobility of a building is not
required, and they possess instead a wonderful degree of ELASTK ITY.
The rye straw bends before the wind, but only to return to its
original position when the force of the wind has been expended.
The mechanical equipment of plant bodies is peculiar to themselves,
but perfectly adapted to their needs. The firm but at the same time
elastic materialwhich plants produce, is put to the most varied uses
by mankind wood forms an easily worked yet sufficiently durable
;
the
building material, and the bast fibres are used in the manufacture of
thread and cordage.
In young stems and plants, in which the stiff but elastic wood and
sclerenchymatous fibres are not developed, the necessary rigidity
cannot be attained in the same way as in the older and woody stems.
But although the principal component of such young stems is water
(often 90 per cent or more), they maintain a remarkable degree of
rigidity and elasticity through the elastic tension of their extremely
thin and delicate cell walls.
Turgidity. When air orwater is forced, under pressure, into an
elastic receptable such as a rubber tube, the walls of the tube become
SECT, ii PHYSIOLOGY 179

stretched and the tube longer and thicker. By this process the tube
becomes just so much stiffer and firmer the greater the internal pressure
and the more elastic and thinner its wall. By the similar tension of
their elastic cell walls arising from internal pressure, the rigidity and

elasticity of thin-walled plant cells, and organs composed of them, are

maintained. The internal pressure results from the osmotic attraction
and interchanges between the substances dissolved in the cell sap and
the water outside the cell.
When twosubstances in solution are separated by a membrane
which permeable to both, a mixture of the two solutions will take
is

place through the dividing wall. If, however, the membrane is more

easily permeated by one of the solutions than by the other, then

a larger quantity of the one than the other will pass through it.
If the membrane is
semipermeable, i.e. is
only permeable for one
solution, that one alone will be drawn through it. If a pig's bladder
be filled with a solution of common salt and then immersed in water,
the flow of water into the bladder is more rapid than the outflow of
the salt solution, and in consequence, an internal pressure is exerted
within the bladder sufficient to expand it to a hard rigid body.
A
pressure similar to that arising from the osmotic attraction of
the salt solution is produced in plant cells by the substances, particu-
larly organic and inorganic acids, salts, and sugars, held in solution in
the cell sap. The living protoplasm of the cell does not allow any of
the substances dissolved in the sap to pass out except such as escape
through the diffusion taking place between the cells themselves, in the
constant translocation and exchange of substances within the plant. As
may be directly observed in cells with coloured cell sap, the dissolved
substances are held in by the protoplasm. These substances, however,
draw in water through the cell walls and the protoplasm, and so set
up a pressure within the cells of at least 3*5 atmospheres, and often as
great as 5-10 atmospheres. In some instances this pressure may
amount to 15, 20, or more atmospheres. In the cambial cells of trees
a pressure of 25 atmospheres, and in the "nodes" of grasses, of 40
atmospheres is attained. Thus a tension is created which frequently
exceeds that exerted by the steam of the most powerful locomotives.
Through the force of such a tension the cell walls become so dis-
tended, that cells under the influence of this internal pressure or
TURGIDITY become longer and larger than in their unexpanded
condition.
When, from any cause, the quantity of water in such a turgescent
cell is diminished the internal pressure is naturally decreased, and the
cell walls, the distension of which may have amounted to 10-20
per cent, again contract. The cell grows smaller, and, at the same
time losing its rigidity and elasticity, becomes soft and flaccid.
This condition occurs from natural causes when a succulent plant
loses more water by evaporation than it can replace, and, in
180 BOTANY

consequence, wilts. Such a flaccid plant plainly shows that the

rigidityis not maintained
by its framework of cell walls, but by the
hydrostatic pressure within the cells, for with a more abundant water-
supply it returns to its original condition.
In addition to loss by evaporation, water can also be withdrawn
from cells by the same molecular force which causes the internal or
endosmotic pressure. In cases where the cells
are surrounded by a solution which exerts an
attraction upon water, the turgidity of the
cells is proportionally weakened, and may be

altogether overcome. On account of the con-

sequent PLASMOLYSis, or the contraction and
separation of the protoplasm from the cell walls,
occasioned by the withdrawal of water, the
tension of the cell walls is decreased, and the
cell becomes flaccid and collapses (Fig. 174).

Solutions of non-metallic organic substances possess

the same osmotic power when the number of molecules
dissolved in the litre is the same their " equimolecular ;

solutions" are "isosmotic." On account of their dis-

sociationwhen in solution into ions, metallic com-
pounds exert H, 2, 2 times this osmotic effect. These
numbers represent the "isosmotic coefficients" of these
substances.
If placed in pure water, however, the previous tur-

gescence of the cells can be restored, if their proto-

plasm has not been too strongly affected by the action
of the solution. If the protoplasm has been killed
in the process, it becomes permeable to water, and it
FIG. 174. Internodal cell of
jg no ] on g er possible to set up an internal pressure.
Xitella.F, Fresh and tur- . ,. i j c
Fresh sectlons ot Beet placed in water, give up none of
p, with turgor
.
gescent ;

reduced, flaccid, shorter their sugar or colouring matter but after the protoplasm
;

and smaller, the proto- has been killed, the sugar and colouring matter at once
plasm separated from the escape into the surrounding water, and the sections lose
cell walls in folds; .<w, ,, .
c , ,.,
their firmness and rijndity.
J
lateral segments, (x circa
6 \ On the other hand, r ungi, Bacteria, or marine plants,
when placed in a weaker saline solution or in fresh water,
have the internal pressure of their cells increased. This may even lead to the

rupture of the cell wall (plasmoptysis).

Through a knowledge of the strength of a solution necessary to produce
internal pressure within plant
plasmolysis, a means is afforded of measuring the
cells. For example, if a solution of saltpetre with an osmotic pressure of 5
atmospheres (a 1 per cent solution, according to PFKFFKK'S investigations, gives
rise to a pressure of about 3^ atmospheres) is just sufficient to overcome the

turgidity of a plant cell (which in the case of stretched elastic cells shows itself by
the limit of contraction being reached), then, conversely, the cell sap exerts
upon water an equivalent endosmotic force required to forcibly
pressure. The
stretch a flaccid or plasmolysed organ to its original length furnishes also a rough
:t
means of estimating the pressure developed in turgescent tissues ( ).
 PHYSIOLOGY 181

In the tension produced by turgidity we see how purely physical

processes determine the rigidity of plants. These PHYSICAL processes
are, however, dependent upon the VITAL functions of plants, inasmuch
as they can only be called into action by living protoplasm in
conjunction with the structure and properties of the cell. Living
plant cells have thus power to regulate the physical effects of
osmotic pressure by increasing or diminishing, or even suddenly
overcoming their turgidity. (The osmotic pressure is increased three
times by the change of glucose into oxalic acid.) It will also be

apparent, in considering the operation of other physical forces, that

the primary and essential result of the vital action is to give rise to
the operation of physical processes, and to favour, constrain, or vary
them in such a way that they become of service to plant life.
Tension of Tissues. The rigidity of parenchymatous tissue,
although to a large extent dependent upon the tension arising from
the turgidity of its individual cells, is nevertheless considerably
enhanced by the opposing pressure between the inner and outer
tissue systems, in particular, between the pith and the epidermal
and cortical tissues. The pith in this case plays the same part as the
cell sap, since it is continually striving to increase its volume ; the

epidermal and cortical layers, on the other hand, by the pressure of

the internal pith cylinders, are stretched and distended, just as are
the cell walls by the osmotic pressure of the cell sap.
The tension of tissues is easily demonstrated by removing a strip
of the peripheral tissue from a piece of a turgescent stem (of a Sun-
flower, Helianthus, for example), and cutting out the pith. It will be
found that the outer tissue at once becomes shorter, and the pith
longer than when they were united together in the stem. If
the length of the stem experimented upon was 50 cm., the cortical
strip would shrink to 46 cm., and the pith suddenly lengthen
to 60-70 cm.

According to J. C. MULLER a pressure of 13 atmospheres would be needed to

prevent the expansion of the isolated pith.
From this experiment it will be seen that the natural length of a stem represents
the equilibrium maintained between the tendency of the pith to elongate and of the
outer tissues to contract. The cortical tissue between the epidermis and the pith
affords a transition between the two extremes of tension, the inner cell layers are

compressed like the pith, and the outer layers stretched like the epidermis. The
tension of tissues is also demonstrated by the fact that each strip of a fresh shoot
which has been split longitudinally will curve outward, so that the pith forms the
convex, the epidermis the concave side.
There is often a great difference in tension even between the outer and inner
layers of the tissue of hollow organs, such as the stalks of a Dandelion ( Taraxacum
officinale), which, when split longitudinally, curl into helices of many turns,
especially if placed in water. A tension exists wherever resistant and unequally
strained tissues are in contact, and often occurs in parts of plants where it does not
182 BOTANY PART i

assist, as inthe leaves and stems, in maintaining the rigidity of the plant body.
Longitudinal and transverse tensions occur, particularly when, through secondary
growth, newly formed growing tissues have to overcome the resistance of other
tissues. In this way the primary and later the secondary cortex of trees become
greatly stretched by the new cambial growth, so much so, that if a ring of bark
be removed from a stem and then placed round it, a force of ten atmospheres is
needed to make the edges meet ;
this was shown by an experiment of KUABBK.
In the meristematic tissues of growing points there is scarcely any perceptible
tension, while, on the other hand, in regions which are in a state of elongation the
tension of the tissues attains its highest limit. After an organ has completed its
growth the elasticity of the cell walls and the turgoscence of the cells decrease ;
and
the tension of the tissues is therefore also diminished. The requisite rigidity is,

however, now provided for special groups of cells with thickened and hardened
by
walls, which thus constitute a firm framework for the other tissues similar to the
bony skeleton of the higher animals.

Mechanical Tissues (Stereome) ( 4 ). The supporting framework

of plants is
provided by the thick-walled elements of the wood, the
thickened sclerenchymatous fibres of the fundamental tissue and the
bast, and more rarely by groups of stone-cells. The firm thick walls
of these tissues are not infrequently further hardened by deposits of
mineral substances. The resistance which these forms of tissue offer
when the attempt is made to cut, tear, or break them affords
sufficient evidence of their hardness, tenacity, and rigidity. More-
over, SCHWENDENER has been able to determine their mechanical
value by means of exact physical experiments and investigations.
According to such estimates, the sustaining strength of sclerenchy-
matous fibres is, within the limits of their elasticity, in general equal to
the best wrought iron or hammered steel, while at the same time
their ductility is ten or fifteen times as great as that of iron.
It is true that soon after exceeding its limit of elasticity the
stereome of the plant becomes ruptured, while the modulus of
rigidity for iron is not reached until the load is increased threefold.
It is, however, of value for the needs of the plant that its limit of

elasticity extends almost to the limit of its rigidity.

Just as the mechanical tissues of the internal framework of plants
exhibit the physical properties most essential for their purpose, their
arrangement, as SCHWENDENER showed, will also be found equally
well adapted to the various ends in view, according as they may be
required to withstand the strain of flexure, traction, or pressure. To
withstand bending, and to offer the utmost possible resistance to
it, a peripheral disposition of the rigid mechanical tissue is the most

favourable. When a straight rod (Fig. 175) is bent, the convex

side elongates and the concave side contracts, that is, the outer

edges (a, a and a', a) are exposed to the greatest variations in length,
while, nearer the centre (i, i and i', i') the deflection and consequent
variations in length are less. Accordingly, if the supporting skeleton
of a plant stem be placed near the centre (i, i'), then
a considerable
SECT. II PHYSIOLOGY 183

degree of curvature is possible with but little flexure of the

mechanical tissue. Nearer the periphery it would be subject to
greater strain, and so offer a greater resistance to the deflecting

/ I

l'n;. 175. Longitudinal section of an elastic cylinder, before bending (dotted out-
I.

line) and after bending (heavy outline). After bending the convex side (a') is
stretched and the concave side (a) compressed. F, connecting tissue.
2. When the connecting tissue (/) is not sufficiently firm, the bands of stereome
(a, a') curve independently and remain unaltered in length.

force. It is essential that the girder- like arrangement of the

stereome should be held together by a connecting tissue which is at
once resistant and elastic. In the absence of such connecting tissue,
the place of which in hollow members may, however, be taken by

SoR
\

. liii. Rigidity against bending. 1. Transverse section of a young twig of SambiKiis; c, collen-

chymn. 'art of the transverse section of a haulm of grass (Mullnia enerulea); sc, ribs of
-'. 1

sclerenchyma sc R, sclerenchymatous ring connecting them laterally

; A, green assimilatory
;

tissue ; MH, pith-cavity.

firm tangential connection of the mechanical tissues, the individual

ribs of stereome are easily bent (Fig. 175, 2). In erect stems and
flower-stalkswhere rigidity is an essential requirement, the mechanical
tissue situated at the periphery, and often takes the form of
is

projecting ridges (Fig. 176, 1, 2).

184

In roots, and in many rhizomes and stolons, which must push

between impeding obstacles, the skeletal system is central, where
without interfering with the bending, it can more effectually sustain
strains upon the longitudinal rigidity of the organ by being thus
massed together (Fig.
f> 177). The stereome is

more highly developed

in those roots which
mainly serve to attach the
plant to the substratum
than in those the main
duty of which is to absorb
r
Fin. 177. Mechanical tissue of roots. 1, Centrally placed
food material ( ').
to resist longitudinal pulling .strains ; 2, a prop root with
Where, however, pres-
a peripheral layer of mechanical tissue (P)to resist lateral
sure must be guarded
pressure, in addition to the central strand.
against (as in Plum-stones,
and in Hazel-nuts and Walnuts), and also
in some subterranean or

submerged parts of plants, the mechanical resistance is maintained by

an arching mass of sclerotic cells or sclerenchymatous fibres.
Stems of trees which have to support heavy and frequently large
crowns, must, like pillars, be constructed to withstand the longitudinal
pressure as well as the bending.
Leaf-blades must be strengthened not only against bending forces,
but against tearing of their expanded surface by the action of wind

FIG. 178. Leaf of Phonniumtenax. 1. Transverse section Sc, plates and strands of sclerenchyma
; ;

Aj green assimilatory parenchyma H, hypoderma serving for water-storage

; ;W, colourless
mesophyll (internal water-storage tissue). 2. Edge of the same leaf E, thick brown epidermis
; ;

7,', marginal strand of sclerenchyma fibres.

or currents of water. The former requirement is met by the vascular

bundles which often project as ribs on the lower surface, and by
girder-like arrangements of the stereome (Phormium tenax, Fig. 178, 1).
Protection against tearing forces is mainly afforded by the firmness
of the margin of the leaf this is partly attained by the vascular
;

bundles forming marginal arches or running parallel to the margin,

in part by special marginal strands of stereome (Fig. 178, 2). Where,
as in the Banana, both these means of protection are wanting, the
lamina is regularly and easily torn by the wind.
 PHYSIOLOGY 185

Collenehyma. All such strongly thickened, inflexible skeletal

elements have lost their capacity for .

growth, and cannot, therefore, be _M^___^ ^ ^/

utilised in those parts of plants which
are in an actively growing state. In
such cases when further support is re-
quired than is afforded by turgidity and
tissue tension, it is secured by the de-
velopment of COLLENCHYMA (p. 68). In
spite of the amount of water it contains,
Fio. 179. Diagrammatic transverse sec-
this tissue, according to AMBRONN'S re
show method of securing
tions, to
searches, opposes considerable resistance rigidity by folding and rolling. In
to tearing, while it is easily stretched the outspread organ A, to withstand
a pressure acting in the direction of
by a longitudinal pull. When the part the aiTow, only the thickness d is
of the plant containing it is growing more available, but in the folded organ B
or less actively, the collenchyma thus the thickness D', and in the rolled
organ <' the thickness D".
readily accommodates itself to the
change in form. It forms, so to speak, the cartilaginous tissue of
6
plants ( ).

Fio. ISO. Buttress roots of a species of MC/VM//'". (After HAHERI.AXDT: from SCHIMPER'S
Pla/nt-Geography.)

Since, as has already been pointed out, the resistance of the

186 BOTANY

mechanical elements to flexure is greater the farther they are removed

from the centre of an organ, it will be readily seen that, while a
flattened, outspread organ can be easily bent, its power of resisting
a deflecting force would be increased if it were folded or rolled
together. In accordance with
this principle many leaves become

plaited or rolled (Fig. 179), and

so acquire a sufficient rigidity
without the assistance of any
specially developed mechanical
tissues.
In addition to the rigidity
of the individual organs of the
plant, their arrangement and
position are of importance for
the stability of the whole organ-
ism. The lateral branches,
which are normally arranged as
symmetrically as possible, may
be forced by bending them up-
wards or downwards to take on
an elliptical cross-section or to
form knee-like bends. In Coni-
"
fers the white wood," which
offers greater resistance tostretch-
FIG.isi.-Young plant of Lupine, the main root of
" red
which has become curved. The lateral roots
ing forces, and the WOod,"
have arisen on the convex faces of the curves,
and act as stays.
wh j ch fe d offerg reate ,.
. .

resistance to compression, ;ire

formed on the sides which are respectively stretched or compressed (").

The roots of tropical trees are so excentrically constructed as to
many
form relatively thin plank-like buttresses to the trunk (Fig. 180).
Lateral roots originate from the convex flanks of a main root
which has been thrown into curves (Fig. 181), and act as stays,
preventing the straightening of the root and the loosening of the hold
of the root-system which would result from this ( 8 ).

II. Nutrition

By all the processes of METABOLISM, or

nutrition are understood
the chemical transformation and conversion of matter carried on
by plants in the production and appropriation of their food supply.
Without nourishment and without the supply of new formative
material neither growth nor development is possible, nor, indeed,
without continuous nutrition can a plant maintain itself in any given
stage of its development. The processes of life are connected with
 PHYSIOLOGY 187

constant changes of the living substance, both transformations and

excretions. Even when the supply of food ceases these processes
continue, so that the death of the organism from starvation must
ensue if the substances that are used up are not at least replaced
in the process of nutrition.
The Constituents of the Substance of the Plant. By means of
chemical analysis the constituent substances of plants can be most
accurately ascertained. It requires, however, no analysis to realise
that a part, often indeed the greater part, of the weight of a plant
is derived from the water with which the whole
plant is permeated.
Water not only fills the cavities of living, full-developed cells, but it
is also
present in the protoplasm, cell walls, and all organised
structures. By drying at a temperature of 110- 120 C. all water
may be expelled from vegetable tissues, and the solid matter of
the plant will alone remain. The amount of dried substance in
plants varies according to the nature and variety of the plant and of
the particular organ. In woody parts it constitutes up to 50 per cent
of their weight, but in herbaceous plants amounts to only 20 or 30

per cent. In succulent plants and fruits it makes up only 5-15 per
cent of their total weight in water-plants and Algae, 2-5 per cent,
;

while all the rest is water.

The plants is combustible, and consists of
dried substance of
organic compounds, which contain but little oxygen, and are con-
verted by combustion into simple inorganic compounds, for the most
part into carbonic acid and water. The elements CARBON, HYDROGEN,
and OXYGEN form the chief constituents of the combustible dried
substance. Next to them in quantity is NITROGEN, which is derived
principally from the protoplasm. After combustion of the dried
substance of plants there always remains an incombustible residue,
the ASH, consisting of the mineral substances contained in the plant.
As these mineral substances undergo transformation during the
process of combustion, they are found in the ash in chemical com-
binations different from those in living plants. From numerous
analyses made of the ash of a great variety of plants, it has been
determined that nearly all the elements, even the rarer ones, may
be present in plants.

In addition to the four already named, the elements met with in the ash of
plants are sulphur, phosphorus, chlorine, iodine, bromine, fluorine, selenium,
tellurium, arsenic (which may be combined as a superphosphate in the soil),
antimony, silicon, tin, titanium, boron, potassium, sodium, lithium, rubidium,
calcium, strontium, barium,
magnesium, zinc, copper, silver, mercury, lead,
aluminium, thallium, chromium, manganese, iron, cobalt, and nickel.

Many these elements, indeed, occur only occasionally and

of
accidentally,while others sulphur, phosphorus, chlorine, silicon,
potassium, sodium, calcium, magnesium, and iron are met with in
188 BOTANY

almost every ash. As might be inferred from the irregular occurrence

of of the elements, they are not all necessary for nutrition, and
many
although their occasional presence in a plant may sometimes change
certain of its special character-
istics (thus the presence of zinc

produces the so-called calami ne

varieties, for example, Thlaspi
alpestre var. calaminare, Viola
lutea var. calaminaria (Fig. 182),

etc.), they do not exerciseadeci-

sive influence upon its existence.
The Essential Constituents
of Plant Food. Chemical
analysis, while enabling us to
determine the substances pres-
ent in plants, does not show
how far they are essential for
nutrition. From culture experi-
ments, in which the plants are
grown in a medium of which
the constituents are known, and
kept under chemical control, it
nas ^ een ascertained that, in
addition to carbon, hydrogen,
oxygen, and nitrogen, which
form the principal part of the
combustible elements of the
dry substance of plants, sul-
phur, phosphorus, potassium,
calcium, magnesium, and iron
are absolutely indispensable to
the growth of all green plants.
In the absence of even a single
one of these elements no normal
development is possible.

According to MOLISCII, only nine

FIG. 182. Viola lutea var. calamity, which has
;<>. of these elements are required by the
longer stems, smaller flowers, and is more branched ]? lln gi It i s not
' however ' iron
than the type form. (i nat. size, from SCH.MPKK'S
gh be supposed, but either
Plant-GeograpJiy.)
calcium or magnesium, that is
unessential. On the other hand,
the ten substances named nutrition of most green plants ; but
suffice for the
it is not to be denied that certain other substances are of use in the plant

economy and of advantage to growth, although not indispensable. Thus, for

example, Buckwheat flourishes better when supplied with a chloride, and the
presence of silica is advantageous as contributing to the rigidity of the tissues.
 PHYSIOLOGY 189

It has also been discovered that by the presence of certain substances, in them-
selves of no nutritive value, the absorption of actual nutritive matter is increased.
In minimal doses poisonous substances often have a favourable effect ; they
lead to better utilisation of the substances at the disposal of the plant and
increase the "economic co-efficient."Their effect is, however, often injurious even
when diluted so as to be imperceptible to chemical tests; thus by such "oligo-
dynamic" influence copper sulphate, even in the proportions of 1:25,000,000, has a
fatal effect on Spiroyyra, and on Peas in a water culture 9
( ).

The nutritive substances are, naturally, not taken up by plants

as elements, but in the form of chemical compounds. CARBON, the
essential component of all organic substances, is obtained by all green

plants solely from the carbonic acid of the atmosphere, and is taken
up by the leaves. All the other constituents of the food of plants
are drawn from the soil by the roots. HYDROGEN, together with
OXYGEN, is obtained from water; oxygen is derived also from
the atmosphere and from many salts and oxides. NITROGEN
is taken up by the higher plants only in the form of nitrates or

ammonium salts ; certain Fungi, Algae, and carnivorous plants, how-

ever, obtain it in the form of peptone, amido-acids, amides, or even
urea. As the ammonia of the soil formed by the soil bacteria from
organic decaying matter is transformed by the help of other so-called
nitrifying bacteria into nitrites, and eventually into nitrates, only
the nitrogen combined in the nitrates need be taken into con-
10
sideration ( ).

Bacteria, as contrasted with the higher plants, are particularly characterised by

their attitude towards nitrogen. In addition to the bacteria, which, by their
nitrifying capability, are of service to green plants, there are other soil bacteria
which in presence of organic compounds of carbon set free the nitrogen of nitro-

genous compounds, and thus render it unserviceable for the nutrition of green

plants. On the other hand, other forms of bacteria (e.g. Azotobactcr chroococcum,
Clostridium pasteurianum), other species of the same group of bacteria, and some-
times also mixtures of certain soil bacteria ( u ) convert the free nitrogen of the air
into compounds whicli serve not only for themselves, but also for other organisms in
the soil and for the higher plants as nitrogenous food material. Whether some
Fungi have the same power is still ah unsettled question. From the comparison
of the crops obtainable from plots of land with and without the addition of
manure, J. KUHN has concluded that a very considerable fixation of nitrogen takes
place in the soil according to ;
WORMBOLD this is in part independent of the action
of micro-organisms ( V2 ).

SULPHUR and PHOSPHORUS form, like nitrogen, important con-

stituents of protoplasm. All proteid substances contain A-2 per cent
of sulphur. The sulphur is taken into plants in the form of
sulphates ; phosphorus in the form of phosphates. POTASSIUM,
unlike sodium, is essential to plant life, and is presumably active in
the processes of assimilation and in the formation of protoplasm ; it
is introduced into plants in the form of salts, and constitutes 3-5

per cent of the weight of their dried substance. MAGNESIUM, like

 190 BOTANY I'AKT I

potassium participating in the most important synthetic processes of

plants, is found in combination with various acids, particularly in
reservoirs of reserve material (in seeds to the extent of 2 per cent)
and in growing points (in leaves only | per cent). CALCIUM also is
taken up in the form of one of its abundant salts, and in considerable

quantities (2-8 per cent). Calcium

plays an important part in the meta-
bolic processes of plants, as a vehicle
for certain other essential substances,
and, as a means of fixing and rendering
harmless hurtful bye -products (oxalic
13
acid) (). IRON, although of the greatest
importance in the formation of chloro-
phyll, is present in plants only in
small quantity.
In order to determine the nutritive
value of different substances the method
of WATER-CULTURE has proved particu-

larly useful (Fig. 183). In these cul-

ture experiments the plants, grown
either directly from the seed or from
cuttings, are cultivated in distilled
water to which have been added certain
nutritive salts. The distilled water
must not have been stored in a copper
vessel, and the salts must be chemically
pure. If all the essential nutritive salts
are present in the culture solution, even
larger plants, such as Indian Corn,
Wheat, Beans, etc., will grow to full
strength and mature seeds as if grown
in earth. It is not necessary in these

experiments to provide carbon com-

pounds in the nutrient solution, as
plants do not derive their carbon supply
FIG. m.-water-cuitures of Fagopyrum through their roots, but, by the help
rscuientitm, i. iii of their leaves, from the carbonic acid
nutrient solution
containing potassium //., in nutrient
;
Q f ^g atmosphere.
solution without potassium. Plants _,, -
,
reduced to sain.- scale. (After XOBBE.) he young plants grow tor a
-I-

time just as well in pure distilled

water as in the nutrient solution but as the supply of nourishment
;

stored in the seeds becomes exhausted, they gradually cease to

grow, and die. If one of the essential constituents of plant food be
omitted from the nutrient solution, although the young plants
will grow better than in the distilled water, they in time become

abnormally developed. When, for example, a plant is

grown in a
 PHYSIOLOGY 191

nutrient solution containing all the essential food elements except

iron, the new leaves developed are no longer green, but are of a pale
"
yellow colour ; they are CHLOROTIC," and not in a condition to
decompose the carbonic acid of the atmosphere and nourish the
plant. Upon the addition, however, of a mere trace of iron to the
solution the chlorotic leaves in a very short time acquire the normal
green colour. The chlorophyll, however, does not contain iron, but
always contains magnesium.
So long as the necessary nutritive substances are provided, the form in which
they are ottered to the plants, as well as the proportionate strength of the nutrient
solution (if not too concentrated), may vary. Plants have the power to take up
these substances in very different combinations, and are able to absorb them in
other proportions than those in which they occur in the soil. In concentrated
nutrient solutions the absorption of water is increased conversely, in very dilute
;

solutions it is the salts that are chiefly taken up. The presence also of certain
substances often exerts an active and generally beneficial influence upon the
thus, calcium salts increase the absorp-
capacity for absorbing other substances :

tion of potassium and ammonium salts. Calciphobous plants do not succeed on

soil rich in lime either because the absorption of potassium, magnesium, and iron
is greatly interfered with (Castanea, Sarothamnus), or because the calcium exercises
a poisonous effect (Pinus pinaster Sphagnum, Drosera). Other plants are calcicolous
,

and succeed best with a high percentage of lime in the soil.

In order to avoid the poisonous effects of phosphates and salts of iron, when
supplied in a soluble form, v. r>. CUONE recommends the following nutritive
solution :

Distilled water 1-2 litres.

Potassium nitrate . . . . . 1 '0 gramme.
Ferrous phosphate
Calcium sulphate
Magnesium sulphate
.....
.

.
. .

.
.

.
.

.
0'5
0'25
0'25
,,

,,

.,

A mixture of equal parts of ferrous phosphate and tri-calcic phosphate may be used
in place of the ferrous phosphate in the above formula. The phosphates which are
present as a fine powder in the solution become deposited on the surface of the
roots of plants growing in the fluid. Plants are found to grow better in the above
solution than in those used by KNOOP and SACHS. The growth of Algae is hindered
14
fn this solution ( ).

As a most important result of such culture experiments, it has

been demonstrated that only the ten elements already named are
necessary for the growth of plants ; all other elements, although
present in plants in large quantities, are of subordinate value to
plant life. This is true, for instance, of SODIUM, which, in combina-
tion with CHLORINE, actually predominates in some plants, and
occasions the characteristic development of many of the succulent
salt-plants (halophytes) and also of SILICON, which, as silica, is so
;

abundantly deposited in the cell walls of many plants Equisetaceae,

Grasses, Sedges, Diatoms (in the ash of Wheat-straw 70 per cent,
and of Equisetaceae 70-97 per cent) that, after combustion of their
192 BOTANY

organic substances, it remains as a firm siliceous skeleton, preserving

the structure of the cell walls. The hardness and firmness of the
cell walls are greatly increased by these siliceous deposits; they
themselves have about the de-
gree of hardness 2, but the
epidermis of Equisetum
silicified
has the hardness 4 ( = Fluor-
spar), and that of the fruits of
Coix 7 ( = Quartz). Some of
the Equisetaceae are even used
for polishing and scouring.
The silicified cell walls of
Diatoms occur as fossils, and
form deposits of SILICEOUS
EARTH (Kieselguhr) in some
geological formations. The
value of the siliceous concre-
termed "
tions, tabasheer,"
that are found within the
joints of the Bamboo has not,
as yet, been satisfactorily ex-
plained. ALUMINIUM, although
like silica everywhere present
in the soil, is only in excep-
tional instances taken up by
plants (Lycopodiaceae, Lichens,
Vitaceae, Leguminosae). Some
species of Lycopodium contain
a sufficient quantity of acetate
of aluminium to render the sap
useful as a mordant. The
same salt is found also in
Grapes. On the other hand,
although scarcely a trace of
iodine can be detected by an
analysis of sea -water, -it is

found, nevertheless, in large

quantities in seaweeds, so much
so that at one time they formed
Flo. 184. Salii-ofiiiii herliucec, a. characteristic halo-

phyte. (From SCHIMI-BB'S riant-Geography.) the principal source of our

15
supplies of this substance ( ).

The substances which, as culture experiments show, are not indispensable for
the life of the plant are, however, of use in so far as they can replace for some
purposes (such as the neutralisation of free acids, etc.) essential elements of plant
food. The latter are thus available for the special purposes for which they are
indispensable. Thus K can be partially replaced by Na, and Mg by Ca.
 PHYSIOLOGY 193

It was first asserted by BEKTHOLLET (1803), and afterwards emphasised by

LIEBIG and C. SPRENGEL, that the mineral salts contained in plants, and once
supposed to be products of the vital processes of the plants themselves, were
essential constituents of plant food. Conclusive proof of this important fact was,
however, first obtained by the investigations of WIEGMANN and POLSTOHFF ( ).
16

The actual proportions of the more important ash constituents of some well-
known plants can be seen from the following table of ash analysis by E. WOLFF ( 17 ).
The table also shows exactly what demands those plants make upon the soil, that
is, what substance they take away from it, in addition to the nitrates which do

not appear in the ash.

194 BOTANY I-AUT i

the walls to liquids and gases. Although impervious to solids, the

cell
walls of living cglls are permeated with " imbibed water ; and to
"
cell
this
" "
IMBIBITION WATER in the cell walls, together with the physical
character of the cell walls themselves, are due their flexibility, elasticity,
and extensibility. The permeability of cell walls for imbibition water
is only possible within certain limits, so that they thus retain the
character of solid bodies.

Treated with certain chemical reagents (potassium hydrate, sulphuric acid, etc.)
cell walls become swollen and gelatinous, or even dissolve into a thin mucilaginous
.slime. This change in their character is due to an increase in the amount of their
imbibition water, induced by the action of the chemicals; otherwise, tin- wati-r
imbibed by ordinary cell walls is limited in amount. The walls of woody cells taku
up by imbibition about one-third of their weight the cell walls of some seeds and
;

fruits and of many Algae absorb many times their own volume when dry.

THE CELL WALLS ARE NOT ONLY PERMEABLE TO PURE WATER,

BUT ALSO TO SUBSTANCES IN SOLUTION. This fact, that the cell wall
offers no resistance to the diffusion of crystalloid bodies when in
solution, is of the utmost importance for the nutrition of the
plant :

cell walls, on the other hand, which are scarcely or not at all per-
meable to liquids (cuticularised walls), take no part in the absorption
of plant-nourishment, except in so far as they may still be permeable
to gases.
In order that liquids may enter by osmosis into the living cell,
they must first pass through the protoplasm, and in the first place
through the outer limiting membrane of the protoplast which is in
contact with the cell wall. LIVING PROTOPLASM is not, however, like
the cell walls, equally permeable to all substances in solution, but,
on the contrary, COMPLETELY EXCLUDES CERTAIN SUBSTANCES WHILE
ALLOWING OTHERS TO PASS THROUGH MORE OR LESS READILY. Moir
it is able to change its permeability
over, according to circumstances,
and thus THE OUTER PROTOPLASMIC MEMBRANE HAS THE POWER OF
DECISION whether a substance may or may not effect an entr.up
into the cell. The wall of the vacuole exercises a similar but often
quite distinct power over the passage of substances from the proto-
plasm into the cell sap. The same influence is exercised by these
membranes on the transfer of substances in a reverse direction.
On account of the selection thus exercised by the protoplasm,
the contents of a cell, in spite of continued osmotic pressure,
are often of quite a different chemical nature from the immedi-
ately surrounding medium. To this same peculiar quality of the
protoplasmic membranes is also due the SELECTIVE POWER of cells,
manifested by the fact that different cells, or the roots of different
plants, appropriate from the same soil entirely different compounds ;
so that, for instance, one plant will take up chiefly silica, another
lime, a third common salt. The action of Seaweeds in this respect
 PHYSIOLOGY

is even more remarkable ;

a medium containing 3 per
living in
cent of common salt, and but potassium salts, they neverthe-
little
less accumulate much larger quantities of potassium than sodium.
In addition they store up phosphates, nitrates, and iodine, sub-
stances which are all present in sea-water in such small quantities
as scarcely to be detected by chemical analysis. Penicillium glaucum
is able to
grow on a nutritive solution containing 21 per cent of
sulphate of copper, owing to the power it possesses of allowing the
entrance to the cell of the salts it requires, while preventing that of
the copper sulphate ( 20 ).
That osmosis may continue into a cell, it is essential that the
absorbed material should become transformed into something else,
either by the activity of the protoplasm or by some other means. Local
accumulations of sugar or other soluble reserve material in fruits,
seeds, bulbs, and tubers would otherwise not be possible ; for osmotic
action, if undisturbed, must in the end lead to the uniform distribution
of the diffusible substances equally throughout all the cells. But if
equilibrium is prevented by the transformation of the diffusible sub-
stances into others that are indiff'usible, the osmotic currents towards
the transforming cells will continue, and the altered and no longer
diffusible substances will be accumulated in them. In this manner
glucose passing by osmosis into the cells of tubers or seeds becomes
converted into the insoluble polysaccharide, starch. As a result of
this a constant movement of new glucose is maintained towards these
cells, which thus become reservoirs of accumulated reserve material

Water and Mineral Substances

Without water there can be no life. THE LIVING PORTIONS OF
ALL ORGANISMS ARE PERMEATED WITH WATER; it is Ollly when ill this
condition that their vital processes can be carried on. Protoplasm,
the real physical basis of life, is, when living, of a viscous, thinly
fluid consistency, and when dried either dies or becomes inactive.

Protoplasm, when in a state of inactivity, as in spores and seeds, can often

endure a certain degree of desiccation for a limited time. During such periods its
actual vital functions cease, and only renew their activity when water is again
supplied.
With the exception of some succulent plants which are uninjured by the loss
of nine-tenths of their water, plants as a rule have their activity impaired by the
loss of water iu withering and are killed by complete desiccation. It is always to lie

regarded as due to some special provision or exceptional quality when entire plants
or their reproductive bodies which have been dried can be again brought to life
by a supply of water. Thus, for example, some Algerian species of Isoetes and the
Central American Selaginclla lejtidophylla can withstand droughts of many months'
duration, and on the first rain again burst into life and renew their growth. In
like manner many Mosses, Liverworts, Lichens, and Algae growing on bare rocks,

tree-trunks, etc., seem able to sustain long seasons of drought without injurv.
196 BOTANY I-AKT i

Seeds and spores, after separation from their parent plants, remain productive for
a long time seeds of Nelumbium, which had been kept dry for over one hundred
;

years, proved capable of germination. A similar vitality was shown by moss

spores which had lain in a herbarium fifty years. The often-repeated assertion
concerning the germination of wheat found with Egyptian mummies ("mummy-
wheat") has, however, been shown to be erroneous. Many seeds lose their power
of germination after having been kept dry for only a year others, even after a few
;

days and others again cannot endure drying at all.

;
On the other hand, the
seeds of some water plants (Eichhornia, etc.) germinate better after being dried
for a period. It must not be forgotten that in all these instances a certain
amount (about 9-14 per cent) of hygroscopic water is retained by plants even when
the air is quite dry. Over the sulphuric acid of the desiccator, seeds retain for
weeks 6 per cent or more of their weight of water. Even drying at 110 or the
21
action of absolute alcohol can be borne by some spores and seeds ( ).

Apart from permeating and energising the cells, water has other
and more varied uses in plant life. It is not only indirectly indis-
pensable for the solution and transport of the products of metabolism,
but also directly in that its elements, hydrogen and oxygen, are made
use of in the formation of organic compounds in plant nutrition.
Water thus used (cf. p. 219) may be designated CONSTITUTION WATER ;
for example, in the formation of every 100 grammes of starch or
cellulose 55 grammes of constitution water, and in the formation of
the same amount of glucose 60 grammes of water, are used. Water
is also necessary for the turgidity and consequent rigidity of

parenchymatous cells (p. 179); it is of use in the process of the

growth of plant cells, which take it up in large quantities, and,
through their consequent expansion, enlarge their volume with but
littleexpenditure of organic substance.
Afurther and still more important service which water performs
for plants consists in THE CONVEYANCE AND INTRODUCTION INTO THE
PLANT BODY OF THE NUTRIENT SUBSTANCES OF THE SOIL. Although
a large amount of water is retained in the plant body (up to 96 per
cent in succulent tissiies) for the maintenance of rigidity and enlarge-
ment of the organs, a still larger quantity of the water taken up by
the roots passes through the plant merely as a medium for the trans-
port of nourishment, and is again discharged through the leaves by
evaporation. By this TRANSPIRATION from the aerial part of plants,
the water passing into them from the roots escapes, and at the same
time, by preventing saturation, which would otherwise be produced,
a continuous upward movement of the water is maintained. The
current of water thus produced is accordingly termed the TRANSI-IKA-
TION CURRENT. As the result of evaporation only water, in the form
of vapour, and gases can escape from the plant. As THE WATERY
FLUID ABSORBED BY THE ROOTS CONTAINS SALTS, AND OTHER NON-
VOLATILE SUBSTANCES IN SOLUTION, THESE ON EVAPORATION ARE
LEFT IN THE PLANT AND GRADUALLY INCREASE IN QUANTITY. This
accumulation of mineral salts is absolutely necessary for the plant,
 PHYSIOLOGY 197

for the nutrient water taken up by the roots is so weak in mineral

substances (it contains but little more solid matter than good drinking-
water), that the plant would obtain too little food if it were only able
to take up as much water as it could retain and make use of.
ALL THOSE CONTRIVANCES, IN PLANTS, THEREFORE, WHICH RENDER
POSSIBLE OR PROMOTE EVAPORATION, OPERATE CHIEFLY IN THE SERVICE
OF NUTRITION. Were transpiration not in the highest degree useful
and even necessary for the acquisition of mineral substances, provision
would certainly have been made by plants to restrict it within the
narrowest possible limits. For transpiration increases the amount of
water required by plants disproportionally to their powers of absorp-
tion,and exposes them, moreover, to the danger of perishing through
the insufficiency of their water-supply. In spite of the increased
danger of drying up, as the result of evaporation, special provision is

made by plants for facilitating transpiration (p. 205).

The Absorption of Water. The "water" absorbed by the plant
is not chemically pure water, but a DILUTE WATERY SOLUTION OF
VARIOUS SUBSTANCES FROM THE SOIL AND FROM THE ATMOSPHERE.
Through the peculiar selective power of their cells (p. 194) plants
exercise a choice from among the substances available in the nutrient
solution.
Aquatic plants and the lower land-plants which are but little

differentiated, such as the Mosses, can absorb water by their general

surface. The same is true of many aquatic Phanerogams. These
(Utricularia, Ceratophyllum, Wolffia), like some Hymenophyllaceae of
damp primaeval forests, the epiphytic Tillandsia usneoides and the Sun-
dew, possess as a rule no roots. The roots of many submerged plants,
on the other hand, penetrate the soil and contribute essentially to the
nutrition of the plant.
In plants living on dry land the conditions are quite different ;
their stemsand leaves develop in the air, and they are restricted to
the water held by capillarity in the soil. In order to obtain this
water in sufficient quantities, special organs are necessary, which may
spread themselves out in the soil, and enter into intimate connection
with its particles, in their search for water. These organs must
absorb the water from the soil, and then force it to the aerial
portions of the plant. This office is performed for a land plant by
its root system, which, in addition to
providing the supply of water,
has also the task of mechanically fastening the plant in the soil (~).

Conversely, loose soil is naturally bound together by the branching roots ; and
on this account plants 'have an economic value in holding together loose earth,
particularly on dykes and land subject to inundation.

The small clod of earth, that is as a rule at the disposal of a plant

in its natural habitat, is utilised to the full
by the highly developed
root system, which behaves in a wonderfully purposive manner.
198 BOTANY

ll' the development of the root system of a germinating Beau or Oak be observed,
it will be found that the growing root of the embryo at once penetrates the soil and
pushes straight downwards. Lateral roots
I

are then given off from the main axis, and,

growing either horizontally or diagonally
downwards, penetrate the earth in tin-
neighbourhood of the primary root. These
lateral secondary roots in turn develop
other roots, which radiate in all directions
Fio. 185. Tip of a ruot-liair witli
ides of soil, (x circa -J40.) from them, and so occupy and utilise the
I>art
entire soil at their disposal. The branch-

ing of the root system can proceed in this manner until, within the whole region
occupied by the roots of a large plant, there is not a single cubic centimetre of earth
which is not penetrated and exhausted by them. The
system of branches of one root system often has an
astonishingly great total length, and may amount to
a kilometre in an annual cereal, and to several kilo-
metres in a well-developed Cueurbita.
All plants do not form a deep-growing tap-root like
that of the Oak, Silver Fir, Beet, Lucerne, etc. some ;

confine themselves to utilising the superficial layers

of the soil by means of a thickly branched lateral root
system (Pine, Cereals). The agriculturist and forester
must, accordingly, take into consideration the mode
of branching and growth of the roots of a plant just
as much as the habit of growtli of its aerial portions.
Plants which make use of different layers of soil 'may
be safely cultivated together in the same soil, and
succeed one another in the same ground. For similar
reasons, in setting out trees along the borders of fields,
the deep-rooted Elm should be preferred to the Poplar.
whose roots spread out near the surface.
Desert or xerophilous plants, according to the ob-
servations of YOLKENS, send out deeply penetrating
roots, which only branch profusely on reaching depths
where they find water. When grown in moderately
dry soil the growth of the roots of oats is greater than
when the soil is damp.

In order to secure a still more intimate

contact with the particles of the soil, there
are produced from the surface of roots small,
exceedingly numerous, and fine cylindrical
FIG. 180. Seedling of orpimu
i

bodies, which penetrate the smallest inter- Betulus. r, Zone of root-

stices of the soil, and fasten themselves so hairs near root-tip ; h, liypo-

hw, main root

closely to its smallest particles as to seem
cotyl ; : .-or.

lateral roots; I, I, lent: >.

actually grown to them (Fig. 185). These

epicotyl ; r, cotyledons.
ultimate branches of the root system, which
discover the very smallest quantity of moisture, and seek out the
most concealed crevices in the search for nourishment, are the ROOT-
 PHYSIOLOGY 199

HAIRS, delicate tubular outgrowths of the epidermal cells. Although

they have the diameter of only a medium-sized cell, and appear to

the naked eye as fine, scarcely visible, glistening lines, they often
attain a length of several millimetres and greatly enlarge the ab-
sorbing surface of their parent root. According to F. SCHWARZ the
epidermal surface of the piliferous zone of the roots of Pisum, which has
230 root-hairs to the square millimetre, is thus increased twelvefold.
The root-hairs cover only a comparatively small zone, a short
distance above the growing root-tip. Soon after they have attained
their greatest length, and have come into the closest contact with the

particles of the soil, they die off. Above this advancing zone of hairs
the epidermis of the root becomes again completely divested of root-
hairs (Fig. 186). The older parts of roots take no share in the process
of absorption. They envelop themselves with cork, increase their con-
ducting elements by growth in thickness, and function exclusively in
the transfer of the water absorbed by the younger portion of the
roots. Even in the young roots the absorption seems principally con-
fined to the regions covered with root-hairs, or, in case no root-hairs
are developed, to a corresponding zone of the root epidermis.
Through the intimate union of the youngest roots with the soil,
they are able to withdraw the minute quantity of water still adhering
to the particles of earth, even after it appears perfectly dry to the
sight and touch. There still remains, however, a certain percentage
of water, held fast in the soil, which the roots are not able to absorb.
Thus, SACHS found that the water left by a Tobacco plant, and which
it could not absorb, amounted in cultivated soil to 12 per cent, in

loam to 8 per cent, and in coarse sand to li pe.r cent. Plants may
even obtain a certain quantity of water from soil which is frozen
hard or from a block of ice.

The ABSORPTIVE POWEU of soil depends, partly, upon chemical changes taking
place within it, but partly also on physical processes (the superficial adhesive
force of its particles). The chemical changes mainly take place in soils rich in clay,
lime, or humus, and containing double silicates of alumina. Salts of potassium
and ammonium, also those of calcium and magnesium, and phosphates are absorbed
by these soils. The former form silicates or double silicates that are only with
diliiculty dissolved, while the phosphoric acid is held combined with calcium or
iron. Magnesium and calcium salts are in other soils but slightly absorbed.
They are, like the chlorides, the nitrates, and, in part, also the sulphates, easily

displaced ;
in soil treated with a solution of saltpetre, for example, the potassium
will remain in combination in the soil, while calcium nitrate passes off in solution.
Humus acids contribute, to a certain extent, to the chemical changes occurring
in soil, as do also soil bacteria, which possess strongly oxidising and reducing
powers (cf. p. 189).
The absorptivity of the soil, which, moreover, is not absolute, and varies with
different soils (sandy soil absorbs poorly), operates advantageously for plants by
the consequent rapid accumulation of large supplies of food -material for their
gradual absorption.
200 BOTANY

The absorptive power of soil for water is due to its capacity to retain water by
capillarity, so that it does not drain oft". Of the soils investigated by S.\< us,
cultivated soil retained in this way 46 per cent, loam 52 per cent, and sand only
21 per cent of water.

The young roots, and especially the root -hairs, in addition

to the carbonic acid exhaled by them, which, no doubt, aids in
loosening the soil, excrete other acid
substances which are able to dissolve
about the same amount from the soil as is
done by a 1 per cent solution of citric acid.
Roots growing upon a polished plate of
marble, dolomite, osteolith, or even ivory
will so corrode them that an etched pattern
of the course and direction of the roots is

thus obtained. Plants which excrete

acid actively (maize, rye, oats) make less
demands on a good worked soil than
those in which the excretion is less active
(wheat, barley) or almost absent (meadow
grasses).
The nutrient water with which the
cell walls of the epidermal cells and root-
hairsfirst become permeated is taken
up by the epidermal cells, and thence
passes through the cortical cells and the
endodermis to the central cylinder of the
root.
The Distribution of the Nutrient
Water. 1. ROOT-PRESSURE.- The causes
ic. is:. vigorous exudation of water which determine the direction and strength
as the. result of root-pressure from
Q{ the movement Q f the water through
a cut stem of DcMia mnabuu. . ,. .
,, ..
,

The smoothly cut stem joined is the living cells of the root-cortex into the
to the glass tube g by means of vascular bundles are not vet fully under-
1

stood. The
fact that the water does
W, absorbed by the roots from the
soil, is pumped out of the vessels actually pass into them, and at times
of the stem with a force sufficient indeed is forced into them with a con-
to overcome the resistance of the
column of mercury Q.
siderable pressure, may be easily de-
monstrated. If the stem of a strongly
growing plant be cut off close above the ground, and the cut
surface dried and then examined with a magnify ing-glass, water will,
in a short time, be seen to exude from the severed ends of the
bundles. By close inspection, it is also possible to determine that
the water escapes solely from the vessels and tracheides of the
bundles. When the soil is kept warm and moist the outflow will be
greater, and will often continue for several days and even months.
The excreted water may amount to several litres (to 1 litre in the
 PHYSIOLOGY 201

Vine, 5 litres in the Birch, and 10-15 (50) litres in the Palms).
This water, as analysis shows, is not pure, but leaves on evaporation
a residue of inorganic and organic substances.

If a long glass tube be placed on the root-stump and tightly fastened by

rubber tubing, the exuded fluid will be forced up the glass tube to a considerable
height. How great the force of this pressure is may be shown by attaching to
the stump a manometer (Fig. 187). The column of mercury will in some cases
be forced to a height of 50 or 60, and under favourable conditions to 100 or more
centimetres (140 cm. in the Birch). These pressures would be sufficient to raise a
column of water 6, 8, and 18 metres high. The height to which the fluid can be
raised is the less surprising when the much greater forces due to turgescence,
which are at the disposal of living cells, is remembered C23 ).
If, instead of the effects of the pressure, the volume of water exuded each hour
be observed, the remarkable fact will be demonstrated that the roots regularly
discharge more water at certain hours than at others (PERIODICITY OF ROOT-
PUESSURE). The quantity is greater by night than by day.
When was shown that the roots were capable of exercising so great a
it

pressure, it was at first believed that the ascent of the sap to the tops of the
highest trees was due to root-pressure. This, however, appears impossible in
view of the following considerations. The volume of water supplied by root-
pressure is not sufficient to replace the quantity given off by evaporation. On the
contrary, during moderately vigorous transpiration, such as takes place on a summer
day, the root-pressure is of a negative character. Thus, if an actively transpiring
plant be cut off near the root, no outflow of water will take place. On the other
hand, the stump will energetically draw in water supplied to it and not until it
;

has become saturated does the force of the root-pressure make itself apparent. In
plants growing under natural conditions, the root- pressure is only effective on
damp, cool days, or at nights, when the transpiration is greatly diminished. In
when the roots are beginning their activity, the conditions are most favour-
spring,
wood is full of water, and the transpiring leaves are not yet unfolded.
able, the
When the wood is injured at this season " sap" is exuded in drops from the vessels
and tracheides.

The so-called BLEEDING from wounds or cut stems is chiefly due

to root-pressure, but it is also augmented by the pressure exerted by
the living cells of the wood (wood parenchyma, medullary rays). FOR
THE LIVING CELLS OF ALL OTHER PARTS OF THE PLANT LIKE THOSE
OF THE ROOT ARE UNDER CERTAIN CONDITIONS ABLE TO FORCE
OUT FLUID. In addition to an excess of water in the plant the
phenomenon may be determined or increased by the stimulus of
wounding or by the healing processes this was shown by MOLISCH for
;

the inflorescences of Palms and in borings made in our native trees.

The amount of fluid excreted from the callus tissue in the latter case
was small, but was forced out even when the pressure amounted to
9 atmospheres. In Schizolobium excelsum Figdor found, in Java, a
24
pressure of 8 atmospheres ( ). The outflowing sap often contains,
in addition to numerous salts, considerable quantities of organic
substances (dissolved albuminous matter, asparagin, acids, and
especially carbohydrates). The amount of saccharine matter in the
_>02 BOTANY

sap of some plants is so great that sugar may be profitably derived

from it. The sap of the North American sugar maple, for example,
contains i per cent of sugar, and a single tree will yield 2-3 kilos.
The sap of certain plants is also fermented and used as an intoxicating
ilrink (birch wine, palm wine, pulque, a Mexican beverage made
from the sap of Agave, etc.). One inflorescence of Agave will yield
1000 litres of sap in from four to five months.

The bleeding which takes place on warm, sunny winter days from wounds o:
in trees is not duo to root-pressure, but to purely physical causes. It is

brought about by the expansion of the

air- bubbles in the tracheal elements i,f

the wood, and be artificially pro-

may
duced at any time in winter by warming
a freshly cut piece of wood when tin- ;

wood is allowed to cool, the air contracts

iin(i t ne water in contact with the cut
e .,, , ,, ,
surface will be again absorbed.
In some lianes the vessels are so wide
that they do not retain the water by
capillarity.When such stems are cut
drinkable water flows from them in con-
siderable quantity.

2. THE COURSE OF THE WATKI:

IN PLANTS. In living plant-tissues
the cells of which require more or
lesswater for their growth and for
the maintenance or augmentation
of their turgidity, there is a con-
stant transfer of water from one
cell to another. This transfer
between the adjacent cells takes
place much too slowly to equalise
the great amount of water lost
188. HAI.I:> experiment to show the ascent
by evaporation from the
ii.

of the sap in the wood. Although the cortex foliage

has been entirely removed at Z, and the wood of a tall tree. IN ORDER TO
:il, me left, the leaves of the branch b remain TRANSKKI! THK WATER, QUICKLY
.'.
as those on the uninjured braneh ,-;
si,

vessel containing water. Facsimile of the

ANJ) m
LAR(JER QUANTITIES, FROM
illustration in HA..F.S' V.,,,,,,1,1, Statta, 17-27.
THE ROOTS TO THE LEAVES, PLANTS
MAKE USE, NOT OF THE LIVINi;
PARENCHYMA, BUT OF THE WOODY PORTION OF THE VASCULAR
BUNDLES. The woody elements which thus conduct the water have
no protoplasm.
3. THE TRANSPIRATION It has long been known thai
CUJIIENT.
the ascending transpiration current in woody plants flows solely
through the wood. It had been observed that plants from which
portions of the cortex had been removed, either purposely or acci-
 PHYSIOLOGY 203

dentally, remained nevertheless perfectly fresh. The adjoining

figure, taken from one of the first books in which the vital processes
in plants were described (ESSAYS ON VEGETABLE STATICS, by STEPHEN

HALES, 1727), shows the method employed in proving this fact experi-
mentally (Fig. 188). At Z in the branch b all the tissues external to
the slender column of. wood have been removed. Since the leaves
of this branch remain as fresh as those of the branch c, it is evident
that the transpiration current must pass through the wood and not
through the cortical tissues. On the other hand, when a short length of
the wood removed from a stem, without at the same time unduly
is

destroying the continuity of the bark, the leaves above the point
of removal will droop as quickly as on a twig cut off from the stem.
It has also been shown by experiment that in herbaceous
plants the
vascular portions of the bundles provide for the conduction of the
ascending currents.

As can be demonstrated by spectroscopical analysis, a dilute solution of li'tliiuni

nitrate taken up by an uninjured plant first ascends in the wood before it passes
laterally into the other tissues. By means of the same solution, MAcNAB, PFITZEI:,
and SACHS determined the velocity of the movement of the transpiration current,
which naturally varies according to the plant and the effect of external conditions
upon transpiration under favourable circumstances it attains a rate of 1-2 metres
;

an hour. The method of showing the exclusive share of the wood in the con-
duction of the water, and, also, of determining the maximum velocity of the
transpiration current, from observations based on the path and rate of movement
of a coloured solution taken up by a plant, is not free from objection, for the

colouring 'matter does not pass through the stem at the same rate as the water
in which is dissolved, but is drawn out and held back by the cells.
it The
employment of coloured solutions will, however, be found instructive for merely
>lio\\ing the course of the transpiration current. The transparent stems of the
Balsam, Imputiens parviflora, and the white- floral leaves of Lilies, Camellias,
ilock Orange, etc., in which the coloured vascular system will stand out as a fine
iirnvork, are especially adapted for such an experiment.

In water plants and succulents, in which little or no transpiration

takes place, the xylem is correspondingly feebly developed. In land
plants, on the other hand, and especially in trees with abundant
foliage, the wood attains a much greater development. All the wood,
however, of a large stem does not take part in the task of water-
conduction, but only the younger, outer rings. Where there is a
distinction between heart- and sap-wood, the heart -wood takes no
part in the conduction of the water.
The character which cause the ascent of the tran-
of the forces
spiration current is
unexplained. Transpiration itself makes a
still

place for the inflowing water. By the removal of the imbibition

water from the cell wall, which is replaced at the expense of the
supply contained in the osmotically active cell, force is exerted which
causes the water in the adjacent elements of the xylem to move
204 BOTANY PAKT i

onwards. It is, however, doubtful whether this initial disturbance

of the condition of equilibrium is sufficient to effect the raising of
the whole mass of water. A sufficient and generally accepted
explanation of this much-debated question is still wanting, though
progress has been made towards it (-').
It has been already explained that the nooT-ruEssuiiE cannot exert such a force
during transpiration (p. 201).
OSMOTIC FORCES act too slowly to be of any value, and, moreover, there is no
fixed distribution of osmotic substances that would account for such a current.
The transpiration current cannot be due to CAPILLARITY. In the first place, con-
tinuous capillaries are entirely wanting in some plants (the Conifers, for example),
and in the stems of others they are only present for comparatively short distances.
Secondly, the concave menisci in the elements of the wood are not in relation with
any level or convex surface of water, in which case alone they could have effect.

Thirdly, the height to which liquids can rise by capillary attraction and it would
be less in the vessels and tracheides than in a glass tube does not approach the
height of an ordinary tree and, finally, the rate of ascent induced by capillarity
;

decreases so greatly with the increasing height of the fluid, that so copious a flow
of water as occurs in plants would be impossible.
ATMOSPHERIC PRESSURE has, also, been shown not to be the cause of the tran-
spiration current. In fact the vessels and tracheides of vigorously transpiring plants
contain rarefied air between the short columns of water. This is evident from the
way which stems cut under mercury become penetrated by it. But as the water-
in
all completely shut oft from the outer atmosphere, the external
courses in plants are
atmospheric pressure could have no effect. The rarefied air within the plants,
moreover, shows no such regularity in its distribution that it could possibly give
continuous a flow of water. Further, as the atmospheric pressure can only
rise to so
sustain the weight of a column of water 10 m. high, while sap rises in Sequoia
100 m. high, and in Eucalyptus trees of the height of 155 m., the inadequacy of
the atmospheric pressure to give rise to such a movement must be .admitted.
The supposition that the water ascends in the form of vapour through the
cavities of the wood, and is afterwards condensed in the leaves, is untenable, as is
at once obvious from a consideration of the anatomical structure of the wood, the

interruption of its cavities by short columns of water, and the temperature of the
plants themselves. And, moreover, the special task of the transpiration current, to
transfer the nutrient salts, could not be accomplished if such were the mode
of ascent.
It has also been suggested that all of these processes might be aided by THE
CO-OPERATION OF THE LIVING CELLS which are so abundant throughout the wood,
and have command of active osmotic forces, to the service of which they could
unite a regulative irritability. STRASBURGER'S investigations, however, have
shown that poisonous solutions, which would at once kill all living protoplasm,
can be transported to the summits of the high trees. Thus the supposition that the
living elements at least immediately co-operate in the ascent of water is precluded.
Recently JOLT, DIXON, and ASKENASY have endeavoured to explain the trans-
mission of the suction force of transpiration to the most distant root-tip by the fact
of the cohesive force of the water. The occurrence of bubbles of air and vapour in
the conducting channels, and the fact that movement of the water interferes with
the power of its cohesive force to resist a pull are among the objections to this theory,
which have, however, been lessened by recent investigations of STIUNHKINK (x ).
 PHYSIOLOGY -205

The recent investigations agree, however, in concluding that THE

TRANSPIRATION CURRENT ASCENDS IN THE CAVITIES OF THE WOOD
THROUGH THE VESSELS AND TRACHEIDES.
4. SUCTION-FOIICE OF TKANSPiiiiSG SHOOTS. A shoot, the cut end of which is

placed in water, shows by remaining fresh that it must be able to draw up water to
itsextreme tips. The force of suction exerted by such a transpiring leafy shoot may
be demonstrated, by fitting the cut end in a long glass tube filled with water in
such a manner that it shall be air-tight. Thus arranged, the shoot will be able to
sustain and raise a column of water 2 metres high. If the lower end of the tube
be inserted in mercury, it will be found that even the heavy mercury will be lifted
by the transpiring shoot to a considerable height. Vigorous coniferous shoots
absorb water through the cut end with a force of suction equal to one atmosphere,
and are thus able to raise the mercury to a height equal to the barometric pressure
(760 mm.), and owing to the cohesion of the water column even beyond this
(920 mm.). The complete exclusion of the external atmosphere is absolutely
requisite for the existence of such a suction-force, and this condition is actually
fulfilled in the water-courses of plants^ 27 ).

The Giving- off of Water. The requisite amount and proper

concentration of the nutrient water supplied by the transpiration
current are maintained only by the constant discharge of the
accumulating water. This may occur in two ways, either by
the evaporation of the water through the cell walls and the
stomata in the form of vapour that is, by transpiration or less
copiously and also less frequently by the actual exudation of drops
of water.
1. TRANSPIRATION. In their outer covering of cork, cuticle, and
wax, plants possess a protection from a too rapid loss of water. A
pumpkin, with its thick cuticle and outer coating of wax, even after
it has been separated from its parent plant for months, suffers no

great loss of water. A potato or an apple is similarly protected by a

thin layer of cork from loss of water by evaporation. The green
organs of plants, o'n the other hand, which must be able to get rid
of the surplus water in order to secure the concentration of the
nutrient salts, make little use of such protective coverings. On the
contrary, they are provided with special contrivances for promoting
evaporation.
The cell walls of all living organs are saturated with water, and,
when the cuticle of the epidermis is not too strongly developed, water
is constantly evaporated, even from uninjured epidermal cells, in
amounts varying with the area of the exposed surfaces. From this
it will be seen that THE FLAT EXPANSION OF FOLIAGE
point of view,
LEAVES RENDERS THEM ADMIRABLY ADAPTED FOR THE WORK OF
TRANSPIRATION. Evaporation is also greatly promoted by the
numerous STOMATA (AIR-PORES) which perforate the epidermis, and
give air, saturated with watery vapour, an opportunity to
the
escape from the intercellular spaces. Although the stomata are so
206 BOTANY i AKT i

small (D'0006 mm. and less) that neither dust nor water can puss
through them into the plant, they are usually present in such
enormous numbers (p. 104) and so suitably distributed that their
united action compensates for their minuteness. When it is taken
into consideration that a medium-sized cabbage leaf (Uraasicd ubT-irm)
is provided with about eleven million, and a Sunflower leaf with
about thirteen million air-pores, it is possible to estimate how greatly
evaporation must be promoted by diffusion through these fine sieve-
like perforations of the epidermis (cf. p. 221).
The stomata also afford plants a means of REGULATING K\ .\P<H;\
TION. The pores, which are the mouths of intercellular spaces, are

Kio. 189. Stoma of lldliliOi us *//. in tran>vrrse set-lion. The ihirkiT lilies show tin- .shape assumed
by the guard-cells when the stoma is open, the lighter lines when the stotna is closed. (After
SCHWENDENER.) The cavities of the guard-cells with the stoma rinsed MV shadeil. .-aid are
distinctly smaller than when the stoma is open.

surrounded by GUARD-CELLS (p. 103). As the term guard-cell sug_

these cells have the power of closing the pore. THE CLOSING \\i>
OPENING OF THE STOMATA ARE ACCOMPLISHED THROUGH CHANGES IN
THE TURGIDITY OF THE GUARD-CELLS OR OF THE ADJOINING El'IDKKM A I.

CELLS. In consequence of their peculiar wall thickenings, elasticity,

and lateral attachment, a change of turgidity affects the size and
shape of the guard-cells in such a way that, on diminished turgidity.
they become flatter and close the air-passage, while an increase of
turgidity has the contrary effect and opens the pore (Fig. 189).

In many plants the so-called subsidiary cells (p. 104) participate in various wa\ .-

and degrees in these processes, depending upon the special structure of the whole

apparatus. The opening and closing of the stomata may follow cither external or
internal influences but such stimuli affect different plants in different manners.
;

Generally speaking, the stomata begin to close on the diminution of the water-
supply, even before this is indicated by wilting they open, on the other hand,
;

when active transpiration is advantageous (in light, in moist air, etc.). The
quantity and quality of the substances held in solution in the nutrient water and
the nature of the surrounding gases react upon the stomata. The size of their
is decreased, and the
opening quantity of water evaporated is therefore ]e.s>> in-.l
when more than the usual amount of nutrient salts is present, in the transpiration
 PHYSIOLOGY 207

current as in that case, if through continued evaporation the nutrient \vat-r

;

should become too concentrated, it might act injuriously upon the plant. In marsh
and water plants the stomata react less promptly than in land plants (**).

It has already been pointed out, in describing the morphology of

the stomata, that they are chiefly to be found on the surfaces of the
leaves. THE LEAVES ARE ACCORDINGLY TO BE CONSIDERED AS SPECIAL
ORGANS OF TRANSPIRA-
TION (and assimilation, p.
214). This is also evident
from the extraordinarily
minute branching and sub-
division of the vascular
bundles in the blade of
the leaf. The adjoining 3*.
illustration 190),
(Fig.
showing the nervature or
distribution of the vascular
bundles in a Crataeyus leaf,
willconvey some idea of
the extensive branching
which the bundles of a leaf
undergo, especially when it
istaken into consideration
that only the macroscopic
and none of the finer
microscopic branchings are
represented in the figure.
By means of this conduct-
ing system, a copious S
supply of nutrient water
can be delivered directly
from the roots to every
square millimetre of the
leaf. There is, however,
a special reason why the
Fio. 190. Course of the vascular bundles (venation) in a
leaves are so abundantly leaf of 'i-ntiii-ijiif.
'
(From a photograph natural size.)
;

supplied. They are the

actual laboratories of plants, in which, out of the carbonic acid of the
atmosphere and the water, and nutrient salts of the soil, the organic
MilllrilKirr Tin o + vi rl r\r fri/i -r^l o VVJ-V^IT -i^ w-w.^-.^J^-.rtrt^
building material of the plant-body is produced.
" TJ^ _ ~
<-
For I
this reason it is
. ," . ,! j.

in the leaves that the broad

expansions of tissue for the special pro-
motion of transpiration are found. The amount of water actually
evaporated from the leaf surfaces in the performance of their vital
functions is surprising. For instance, a
strong plant, Sunflower of
about the height of a man, evaporates in a warm
day over a litre
of water. It has been estimated that an acre of
cabbage plants will
 208 BOTANY PAKT i

give off two million litres of water in four months, and an acre of
hops three to four millions. For a Birch tree with about 200,000
leaves and standing perfectly free, VON HOHNEL estimated that 500
litres of water would be lost by evaporation on a hot dry day ; on an

average the amount would be 60-70 litres. A

hectare of Beech wood
29
gives off on the average about 30,000 litres daily ( ).
It has been calculated that during the period of vegetation the
Beech requires 75 litres and the Pine only 7 litres for every 100
grammes of leaf substance. According to DIETRICH, for every gramme
of dry, solid matter produced, on the average, 250-400 grammes of
"

water are evaporated.

EXPERIMENTAL DEMONSTRATION OF TRANSPIRATION. The evaporation from

plants, although imperceptible to direct observation, may be easily demonstrated,
and its amount determined by the help of a few simple appliances. One method
of doing this, is to weigh a plant before and after a period of vigorous evaporation,
r

and thus determine the amount of water actually lost. Or, if the water evaporated
by a plant placed under an air-tight bell-jar be absorbed by calcium chloride or
concentrated sulphuric acid, it will only be necessary to determine the increase in
weight of the absorbing substance to estimate the amount of water given otf by
evaporation. The amount of water taken up by a plant may also be shown by so
arranging the experiment that the water passes in through a narrow tube, as then
even a small consumption of water will be quickly indicated by the rapid change
of the water-level, which will be the more rapid the smaller the bore of the tube.
An apparatus of this kind is called a potetometer, and by its means the amount
of water taken up by a transpiring branch may be determined.
The important part taken by the stomata in the process of transpiration may be
easily shown, according to STAHL, by means of the cobalt reaction, or the change
in colour of dark-blue dry cobalt chloride to light rose upon absorption of water.
In making this experiment a leaf placed between strips of paper which have been
previously saturated with this cobalt salt and then thoroughly dried, is laid between
glass plates. The paper on the side of the leaf most abundantly supplied with
stomata will then first change its colour, and that too the more rapidly the more
widely open are the stomata. The cobalt reaction, as also the iron and palladium
chloride reaction used by MEUGET, may be used to determine variations in the
width of the stomatal openings. FR. DAUWIN used a delicate hygrometer for this
purpose in order to follow continuously the variations in width of the opening.
By the dimming of a film of collodion through which all the microscopic detail
of the leaf surface could be seen, BUSCALIONI and POLLACCI were able to prove that a
30
slight general transpiration in addition to that through the stomata takes place ( ).
It is evident from these and similar experiments that more water is evaporated
in a given time from some plants than from others. These variations are due to
differences in the area of the evaporating surfaces and to structural peculiarities
(the number and size of the stomata, presence of a cuticle, cork, or hairy covering,
etc.). But even in the same shoot transpiration is not always uniform. This is
attributable to the fact that, both from internal and external causes, not onlv tliu
size of the openings of the stomata varies, but also that transpiration,
just as
evaporation from a surface of water, is dependent upon external conditions. Heat,
as well as the dryness and motion of the air, increases transpiration for purely

physical reasons while light, for physiological reasons, also promotes it.
; From
 PHYSIOLOGY 209

both physical and physiological causes, transpiration is more vigorous during the
day than night. Plants like Impatiens parviflora, which droop on warm days,
become fresh again at thefirst approach of night.

2. EXUDATION OF WATER. The discharge of water in a liquid

state by direct exudation
not of so frequent occurrence as its loss
is

by evaporation in the form of vapour. Early in the morning, after

a warm, damp but rainless night, drops of water may be observed
on the tips and margins of the leaves of many of the plants of
a meadow or garden. These drops gradually increase in size
until they finally fall off
and are again replaced by
smaller drops. These are
not dew-drops, although they
are often mistaken for them ;

on the contrary, these drops

of water exude from the

FIG. 101. 'Exudation of drops of FIG. 192. Resinous covering of the stun of a desert
water from a leaf of Tropaeolum plant (Sareocaulon). (From SCHIMPER'S I'lunt-
mqjus. Geogra.phy.)

leaves themselves. They are discharged near the apex of the leaves
of the Indian Corn, but in the case of Alchemilla from every leaf-
tooth, and of the Nasturtium from the ends of the seven main nerves
(Fig. 191). The drops disappear as the sun becomes higher and the
air warmer and relatively drier, but can be induced artificially if a
glass bell-jar be placed over the plant, or the evaporation in any
way diminished. When
the plant becomes overcharged with water
through the activity of the roots, it is discharged in drops. These
are pressed out of special water-stomata (p. 104) or in other cases
through the ordinary stomata, or from clefts in the epidermis.
1'
210 150TANY I'AUT 1

Sometimes they are secreted by specially constructed hairs, and in

i. I'.'S. llnonllii iniiiiititillni'iK tram w Xnilunil, showing the cushion-like shapi-

of the individual plant. (From SCHIMI'KK'.S l'l<>, .'-',' ,,,/, .^

Datura the water is excreted

through the walls of the ordinary
epidermal cells.
It is possible to cause similar
exudation of water in drops by

forcibly injecting water into a cut

shoot.
Such exudations of water are particu-
larly apparent onmany Aroids, and drops
of water may often be seen to fall within
short intervals, from the tips of the large
leaves. From the leaves of Colocasia
nymjilniffolia the exuded drops of water
are even discharged a short distance, and
190 drops may fall in a minute from
(i
single leaf, while ,V litre may be
secreted in the course of a night. In
Spathodea, a tropical member of the
Bignoniaceae, and some other plants, the
space enclosed by the calyx, in which the
young organs are developed, is filled
floral
with water. Again, in unicellular plants,
especially some Moulds, the copious ex-
udation of water is very evident. The
water in this case is pressed directly
through the cell walls, and in some cases
also, as is the case in water plants, through
31
the easily permeable cuticle ( ).

Fin. i-."4. A< ..... la MarptMota, :m Australian The organs for the discharge
-.rl.'mpiiyllotis plant, showing pl.yllodfs. o f water, which HABERLANDT has
collectively termed .
hydathodes
(p. 108), in some
instances, actively press out the water; or, on the
other hand, they may simply allow it to filter through them when
 PHYSIOLOGY 211

the internal pressure has attained a certain strength, and has

caused the water to fill the intercellular spaces. Hydathodes are
mostly found on young organs and are early developed on them.
Since the excretion of water in the liquid form can occur when
the conditions are unfavourable to transpiration (Lathraea) it may in
a sense take the place of transpiration in maintaining the current
from the water-absorbing organs. Its physiological significance is
not, however, the same as transpiration, since the expressed water

Kirj. 195. Transverse sections of the leaf of ,S/iji i-npiUntn. The leaf above in the
closed state, the half leaf below expanded. U, lower surface, without stomata ;

", upper surface with stomata (,S) ; C, chlorophyllous mesophyll. (x 30. After
KERNER VON MARILAUN.)

always contains salts, and sometimes also organic substances in

solution. In fact the quantity of salts in water thus exuded is often
so abundant that after evaporation a slight incrustation is formed on
the leaves (the lime-scales on the leaves of Saxifrages) ( 32 ).
In some instances, also, the substances in solution in the water
are exuded with a purpose, as in the case of the secretions of
the NECTARIES, of the DIGESTIVE GLANDS of insectivorous plants,
and of the discharge of the viscid STIGMATIC FLUID. The excreted
substances in these instances exert an osmotic attraction on the
water in the cells of the plant ; this distinguishes such cases from
excretion dependent simply on the internal pressure. The substances
excreted by some desert plants (Eeaumuria, Tamarix) are so strongly
hygroscopic that the leaves remain covered with numerous drops of
212 BOTANY I'AIIT I

liquid even in the dry air and under the arid conditions natural to
the plants. The superfluous water is discharged by a few plants, the
Pumpkin, for example, into the
cavities of their stems and leaf-stalks,
and is
again absorbed from these
33
reservoirs when needed ( ).

Special Contrivances for regulating

the Water-supply. Almost all the higher
plants possess, in the power to close their
stomata, a special means of checking
transpiration during a temporary insuflici-
ency of the water-supply. In districts

subject to droughts of weeks' or months'

duration, only such plants can flourish as
are able either to withstand a complete
drying up without injury (p. 195), or to
exist for a long time on a scanty supply of
water (xerophytes). This last case is only
rendered possible by the extreme reduction
of transpiration, or by the formation of
organs in which, in times of a superfluity
FIG. 106. Ve'muca. <v>jj;,
.-. iVr.v, with the habit
of water, it may be retained for later use.
of a Cypress, from New Zealand. '(From
SCHIMPER'S Plant-Geography.) Such protection against excessive
transpiration is afforded by the formation
of cork or cuticular coverings and in exceptional cases coverings of resin (Fig. 192),

by the reduction in the number and size of the stomata, their occurrence in cavities
or depressions, and the more orless complete

filling of the opening by waxy substances.

The rolling up of the leaves, the stomatiferous
surface being on the concave side, as well as
the development of thick growths of hair, or of
a covering of star-shaped or scaly hairs, and the
assumption of a vertical position to avoid
the full rays of the sun, are also measures
frequently adopted to lessen transpiration.
The most efficient protection, however, from
too great a loss of water by transpiration is
undoubtedly obtained by the reduction of the
transpiring surfaces, either through a diminu-
tion in the size of the leaves or through their
complete disappearance. The same result may
be obtained by the crowding of the branches
of the plant to form a dense cushion (Fig.
193).
The upright dured leaves may be in the upper
position of the leaves, or the
globose shoots.
substitution of
expanded, perpendicularly
directed leaf-stalks for the leaves (Pnvi. LOOKS), particularly characterises the
flora of Australia (Fig. 194). A clothing of hair, on the other hand, protects
the leaves of some South African Proteaceae (e.g. Lcucadeiulron argenteum).
Some of the Gramineae (Stipa capillata, Festuca alpestris, Sesleria lenuifolia,
 PHYSIOLOGY 213

,s'.
jinnctoria, etc.) roll or fold their leaf-blades, in times of drought, by means of
special hinge-like devices, into narrow tubes, the protected inner surface being
formed by the side bearing the stomata. Reduction of the leaves is illustrated
by the desert forms of Genista and Sarothamnus, and by the Cypress-like Conifers.
The New Zealand Veronica, shown in Fig. 196, closely resembles these in habit. A
complete disappearance of the whole leaf surface takes place in most Cacti, in which
also the stems become swollen and converted into water-reservoirs. A similar de-
velopment of succulent swollen stems frequently occurs in the Euphorbiaceae (Fig.
197), in the Compositae (Kleinia articulata], Asclepiadaceae, and other plant families
found in arid regions. It has been estimated ( w ) that the amount of water evaporated
by a Melon-Cactus is reduced by its succulent development to g^^ of that given
off by an equally heavy climbing plant (Aristolochia). Instead of the stem the
leaves themselves may become succulent, as in the House-leek and other species of
Sempervivum, also in many species of
Sedum, Aloe, and Ayave. Both stem and
leaves are equally succulent in many
species of Mesembryanthemum. In other
plants, the parenchyma of their stem
tubers (epiphytic Orchids) or of their
thickened roots (Oxalideae) serve as water-
reservoirs. Epiphytic Bromeliaceae catch
the rain-water in reservoirs formed by their
closely approximated leaves, and then
eagerly take it up through the scaly hairs
which cover the leaf surfaces, as in species
of Tillandsia. Again, many epiphytic
Orchids and Aroids collect the rain-water
in a swollen sheath developed from the
epidermis of the aerial root (velamen
j. i/vnv r it. c xi FIG. 198. SaJ no?m.
' Full grown and bear-
radicum, p. 109). In the case of other ,

Jng fmits _ (Nat gjze; from SCHIMPER S

epiphytic Orchids, Aroids, and Ferns Plant-Geography.)
(Asplenium Nidus, for instance), the
humus and other material caught in receptacles formed by the leaves or aerial
roots act like a sponge in taking up and retaining water, while the absorptive roots

penetrate into these moist, compost-like masses and absorb both water and nutrient
substances. Many pecies of Frullania (a Liverwort common on Beech trees)
possess, on the other hand, special water-sacs on the under side of their thallus
(Fig. 386). A particularly remarkable contrivance for maintaining a constant
supply of water is exhibited by the epiphytic Dischidia Mafflesiana, a number of
whose leaves form a deep but small-mouthed urn, into which the roots grow. It
would seem at first sight unnecessary that plants like the Mangrove tree, which
stand with their roots entirely in water, should require protection against too
rapid transpiration ; but, as this tree grows in salt or brackish water, it is necessary,
as in other halophytes (Fig. 184) to reduce the amount of water absorbed, in order
to prevent a too great accumulation of salt in the tissues.
In high latitudes, where the soil remains frozen for months at a time, rendering
the absorption of water by the plant difficult, arrangements similar to those found
in desert plants are present to diminish transpiration. The dwarf habit which is
so frequently characteristic of the alpine and arctic plants (Fig. 198) is as much an

expression of this limitation of the transpiration as of the generally unfavourable

:H
conditions of existence ( ).
214 BOTANY i AI:I i

The Absorption of Carbon (Assimilation)

In any attempt to distinguish the relative importance of substances

utilised in plant nutrition, carbon undoubtedly ranks first. Every
organic substance contains carbon, and there is no other element
which could supply or take part in the formation of so many <>r
such a variety of substances, in living organisms as in the chemical
laboratory. Organic chemistry, in short, is merely the chemistry uf
carbon compounds. Living beings which are composed of organic
substances owe the possibility of their existence primarily to the
properties of carbon.
It requires no chemical analysis to realise that plants actually
contain carbon, although in an imperceptible form. Every burning
splinter of a match shows, by its charring, the presence of this
element. An examination of a piece of charcoal in which the finest
structure of the wood is still distinguishable, shows how abundant is
the carbon and how uniformly distributed. Estimated by weight, the
carbon will be found to make up about half the dry weight (when
freed from water) of the plant. The great quantities of coal in the
deeper strata of rocks are the remains of ancient plants both in
;

peat and in. some cases in coal the macroscopic and microscopic
structure gives proof of this origin.
AVhence do plants derive this carbon? The "humus" theory,
accepted for a long time, assumed that the humus of the soil was the
source of all and that carbon, like all the other nutrient
the supply ;

substances, was taken' up by the

roots. That plants grown in pure
sand free from humus, or in a water-culture, increase in dry substance,
and consequently in carbon, clearly demonstrates the falsity of this
theory. The carbon of plants must therefore be derived from other
sources and, in fact, the carbon in humus is, on the contrary, due to
;

previous vegetable decomposition. The discovery made at the end of

the eighteenth and the beginning of the nineteenth century, that THE
CARBON OF PLANTS IS DERIVED FROM THE CARBONIC ACID OF THE
ATMOSPHERE, and is taken up by the action of the green leaves,
is associated with the names of INGENHOUSS, SENEBIER, THEO. DE
SAUSSURE, and SACHS. This discovery is one of the most im-
portant in the progress of the natural sciences. It was, by no means

easy to prove that the invisible gaseous exchange between a plant

and the atmosphere constitutes the chief source of nourishment ;

and it required the courage of a firm conviction to derive the

thousands of pounds of carbon accumulated in the trees of a forest,
from the small proportion (0'03 per cent) of carbon dioxide contained
in the atmosphere.
 PHYSIOLOGY 215

The amount of carbonic acid gas contained in the air varies at different times
and places. H. BUOWN found that in 10,000 litres of air it was 2'7-2'9 litres in
July, 3 '0-3 '6 litres in the winter ; close to the ground 12-13 litres were present in
the same volume. The average amount is about 3^-3 litres in 10,000 litres of the
atmosphere. This weighs about 7 grammes, of which Tsr is oxygen, and only T y :

carbon. Only 2 grammes of carbon are thus contained in the 10,000 litres of air.
In order therefore for a single tree having a dry weight of 5000 kilos to acquire its
2,500,000 grammes of carbon, it must deprive 12 million cubic metres of air of their
carbonic acid. From the consideration of these figures, it is not strange that the
discovery of INGEXHOUSS was unwillingly accepted, and afterwards rejected and
forgotten. LIEBIG was the first in Germany to again call attention to this discovery,
which to-day is accepted without question. The immensity of the numbers just
cited are not so appalling when one considers that, in spite of the small percentage
of carbonic acid in the atmosphere, the actual supply of this gas is estimated at
about 3000 billion kilos, in which are held 800 billion kilos of carbon. This
amount would be sufficient for the vegetation of the entire earth for a long time,
even the air were not continually receiving new supplies of carbonic acid
if

through the respiration and decomposition of organisms, tli rough the combustion
of wood and coal, and through volcanic activity. An adult will exhale daily about
900 grammes C0 2 (245 grammes C). The 1400 million human beings in the world
would thus give back to the air 1200 million kilos of C0 2
(340 million kilos C).
The C0.2 discharged into the air from all the chimneys on the earth is an enormous
amount. CREDNEK calculated that 460,000,000, 000 kilos of coal are burnt annual]}-,
yielding to the atmosphere about 1,265,000 million kilogrammes of carbonic acid
gas. These sources of C0 2 are, according to Beyerinck, insignificant compared
with the enormous amounts produced by the bacteria of the soil in putrefactive
processes. The whole carbon supply of tbe atmosphere is at the disposal of plants,
since the C0 2 becomes uniformly distributed by constant diffusion.

Not all plants, nor indeed all parts of a plant, are thus able to
abstract the carbon from the carbonic acid of the air. Only such
organs as are coloured green by chlorophyll are capable of exercising
this function, for the chlorophyll bodies themselves are the labora-
tories in which this chemical process, so important for the whole
living world, is carried on. From these laboratories is derived the
whole of the carbon which composes the organic substance of all living
things, plants as well as animals. Animals are unable to derive this
most essential element of their bodies from inorganic sources. They
can only take it up in organic substances, which have been
previously formed in plants. Such plants, also, as are without
chlorophyll, as, for example, the Fungi and some of the higher
parasitic plants, are dependent for their nutrition upon organic
substances previously formed by the chlorophyll bodies of other
plants.
Roots and other organs unprovided with chlorophyll, and also the
colourless protoplasm in the green cells themselves, are similarly
dependent upon the activity of the chloroplasts.
The derivation of carbon from carbonic acid and its conversion
into organic substances is termed ASSIMILATION. In its broadest
216 linTANY PABT I

sense, and especially in the animal kingdom, the word assimilation is

used for nutritive processes by which the nourishment

all is built up
into the substance of an organism. But in Botany the meaning of
the term has gradually been restricted, and now by assimilation the
assimilation of carbon by means of the chlorophyll granules is
specially understood. Moreover, all the further synthetic processes
dependent upon carbon assimilation.
of assimilation are
The chlorophyll
bodies, however, can only produce organic sub-
stances from carbonic acid and water by help of the vibrations of
light (PHOTOSYNTHESIS). The chlorophyll apparatus is unable to
assimilate in darkness, although all the other requirements are
present for active assimilation. Given a source of illumination,
either natural or artificial, assimilation commences, and, within
certain limits, increases in proportion to the intensity of the effective
rays. Unfavourable conditions, such as cold, or the presence of
poisonous substances, may inhibit the action of the chlorophyll
apparatus.
The vibrations of the ether perceptible as light supply the energy
for the decomposition of carbonic acid and the production of carbon,

just as other vibrations, in the form of heat, supply the energy

requisite for the working of a steam-engine. A considerable amount
of work has to be done in the assimilation of carbon, and on the
potential energy thus stored up the vital processes are mainly
dependent. Even the force exerted by the steam engine can be
referred to the work of assimilation which was performed by the
plants whose remains are burnt in the furnace. For in the process of
combustion the organic material is oxidised producing carbon dioxide,
and the energy which was originally needed to produce the substance
of the plants from carbon dioxide is set free.
Not all vibrations are equally capable of arousing the
light
assimilatory activity. Just as the rays of different refrangibility
differ in their action, both upon the eye and the photographic plate,
so they have a different effect upon assimilation. It would be
natural to suppose that the chemically active rays, the blue and
which decompose silver
violet, and other chemical compounds,
salts
would also be the most effective promoting the assimilatory activity
in
of the chlorophyll bodies. Exactly the contrary, however, has been
shown to be the case. The highly refractive chemical rays have
little or no effect on assimilation the red, orange, and yellow rays,
;

that is, the so-called illuminating rays of the spectrum, are on the
3C
contrary the most active ( ).
In the red-leaved varieties of green plants, such as the Purple Beech and Red
Cabbage, the chlorophyll is developed in the same manner as in the green parent
species, but it is hidden from view by a red colouring matter in the epidermis or
in deeper-lying cells. In the Red Algae on the other hand the chromatophores
themselves have a red colour ;
after death this becomes free as a bluish-red pigment
SECT, ii PHYSIOLOGY 217

(phycocyan) leaving the chloroplasts green. The Brown Algae and the Diatomaceae
have a brown modification of chlorophyll (phaeophyll) which after death changes
into chlorophyll.
In the blue-green freshwater Algae, and also in the Brown and Red Seaweeds,
the maximum assimilation takes place, according to ENGELMANK, in another part
of the spectrum than it does in the case of green plants. The assimilation in
these Algae seems indeed to be carried on in the part of the spectrum, the colour
of which is complementary to their own. The pigments associated with the
chlorophyll thus appear to act in the same way as the sensibilisators on the
photographic plate they attract light of a different wave length to co-operate
;

in the chemical process. All the rays of the mixed white light are usually at the
disposal of plants growing freely in the open air ; only the Seaweeds found in deep
water (at the most but 400 m. below the surface) grow in a prevailing blue light,
while the deeper-lying tissues of land plants live in red light, as this penetrates
farther into the parenchymatous tissues ( 37 ).
In studying the effect of different kinds of light upon assimilation, it is custom-
ary either to use the separate colours of the solar spectrum, or to imitate them by
means of coloured glass or coloured solutions. For such experiments it will be
found convenient to make use of double-walled bell-jars filled with a solution of
bichromate of potassium or of ammoniacal copper oxides. Plants grown under jars
filled with the first. solution, which allows only the red, orange, and yellow rays
to pass through, assimilate almost as actively as in white light. Under the jars
containing the second solution, which readily permits the passage of the chemically
active rays, assimilation is much less active.

But little is known with regard to the processes carried on in green cells during
assimilation, and by no means clear what part the green chlorophyll
it is still

pigment performs. The pigment which may be extracted from the protoplasm of
the chlorophyll bodies makes up only a small part of their substance (about O'l per
cent.), and gives no reaction from which its operations may be inferred. The light
absorbed by the chlorophyll pigment also stands in no recognisable relation to the
requirements of assimilation, for the assimilation is not proportional to the intensity
of the absorption of the different rays. The proportion of the energy, passing
through the leaf in the form of light, utilised in assimilation is, according to the
thermo-electric measurements of DETLEFSEN, only 1 per cent according to the
;

calculations of H. BROWN, ^ per cent in sunlight, and over 2 per cent in diffuse
38
light f ). It has not as yet been determined what part the mineral constituents
of the transpiration current take in the process. On the other hand, the proto-
plasmic body of the chloroplasts cannot assimilate when the green pigment is not
present ; that is, when, from any cause, the corpuscles are prevented from turning
green. As the existence of the green pigment is dependent upon the presence of
oxygen, of iron, of carbohydrates and other food substances, upon a proper temper-
ature, and, with few exceptions (Ferns, Conifers, cotyledons of Sycamore, lower
Algae in culture solutions), upon the action of light, its formation in the chloro-
phyll bodies may be prevented by depriving them of the requisites for its develop-
ment. The chromatophores will then remain yellow (in leaves) or white (in steins).
Within recent years it has, indeed, been determined that certain nitrifying
bacteria have the power of forming a small amount of organic substances from
carbonates, carbonic acid, and ammonia. The process by which the organic
carbon compound is derived must, however, be altogether different from that of
green plants, as the bacteria contain no chlorophyll, and their nutritive activity
is in no way dependent upon the light. The necessary energy is here obtained
218 HOT ANY I'AKT I

not from photosynthesis but from the oxidation of ammonia into nitrous acid, and
this into nitric acid (chemosynthesis). The formation of organic substance in tin 1

sulphur bacteria and especially in the so-called purple bacteria is also insufficiently
understood, but photosynthesis appears to play a part in it.

As a result of the chemical processes involved in the decomposing

activity of assimilation, only the special end-product and one bye-
product are at present known. SACHS discovered that the or^nn it-
compound, first to be detected as the special ultimate product of

assimilation in the higher plants, is a CARBOHYDRATE, which may

either remain in solution, or in the form of STARCH GRAINS may
become microscopically visible at the points of its formation. In a
number of plants (e.g. the Algae) the first visible product is often not
starch but a fatty oil, protein, or some other secondary product.
A
short time after assimilation begins,
in sunshine, sometimes within five minutes,
minute starch grains appear in the chloro-
plasts. These grains gradually enlarge until,

finally, they greatly exceed the original

may
size of the chloroplasts. Should, however,
the assimilation cease, which it regularly does
at night, then the starch grains are dissolved
and as soluble carbohydrates (glucose, etc.)
pass out of the cell. In some plants (many
Monocotyledons) there is no starch formed in
the chloroplasts, but the products of assimila-
tion pass in a dissolved state directly into the
FIG. 199. A showing the
leaf :

cellsap. In exceptional cases, however, starch is

iodine reaction. Part of an a ] go f orme(] where there IS a Surplus of glllCOSe,
assimilatini' leaf was covered
with a striper tinfoil. After-
-,
, ,
f
sugar, Jind other substances, as, for example,
m >

wards, when treated with a the guard Cells of the StOTTiata of MoilOCOtvle-
soiution of iodine, the part
dons, and in the coloured plastids of flowers and
of the leaf darkened by the . .

fruits.
T.I r j.-
i
In other cases also only a fraction ot
overlying tinfoil, having
the product of assimilation appears as starch
formed no starch, gave no
colour reaction. nat,
(
in Hdianthus, for example, only J) while the
(}

carbohydrates formed in the first place (sol-

uble hexoses, especially dextrose C 12 6) (!
H
are in part transformed
into di- and poly-saccharides. In a chemical sense the process of
assimilation is thus an asymmetrical one.
The formation of starch may be shown to be a direct result of
assimilation, by means of the "iodine reaction" and without the aid
of a microscope. If a leaf cut from a plant previously kept in the
dark until the starch already formed in the leaves has been removed
in a soluble form, be treated with a solution of iodine after being
first decolorised in hot alcohol, it will in a short time assume a

yellowish-brown colour, while a leaf vigorously assimilating in the

light will, with the same treatment, take a blue-black colour owing
 PHYSIOLOGY 219

to the starch abundantly present in the cells. In Fig. 199 the result
of the iodine reaction is shown on a leaf, part of which had been
covered with a strip of dark paper or tinfoil. The cells darkened by
the overlying paper or foil formed no starch, while those exposed to
the light are shown by the iodine reaction to be full of it. A green
leaf kept in air devoid of carbonic
acid, although fully exposed to the
light, will similarly form no starch.
The bye-product arising from
the assimilatory process is PURE
OXYGEN. The volume of oxygen
thus set free nearly equal to (in
is

most cases it somewhat exceeds)

the volume of carbonic acid
taken in. If plants assimilate in
a known quantity of air containing
carbonic acid gas, its volume will
therefore remain .nearly the same.
The chemical process of assimila-
tion resulting in the decomposition
of the carbonic acid may be thus
expressed :

6C0.2 + 6H.,0 = C 6 H ]2 B + 60.,

(Glucose).

From this chemical equation( 30 )

it is evident that the ELEMENTS OF
WATER ARE REQUISITE FOR THE
PROCESS OF ASSIMILATION (p. 189).
In the transformation of the mono-
saccharides into poly-saccharides
(starch, cellulose) which afterwards ,
Fio. Evolution ot oxygen from assimilating
'JltO.

takes place, small amounts of plants. In the glass cylinder C, filled with
water are again liberated. water, are placed shoots of Elodea canadensi?;

the freshly cut ends of the shoots are intro-

The oxygen given off by green duced into the test-tube B, which is also full
plants, although not perceptible of water. The gas-bubbles E, rising from the
cut surfaces, collect at S. H, stand to sup-
when they are growing in the open
port the test-tube.
air, becomes apparent in the case
of water plants. It was indeed through the evolution of bubbles of

oxygen from water plants that INGENHOUSS first had his attention
called to the assimilatory activity of leaves. To see this process, it
is
only necessary to place a cut stem of a water plant in a vessel of
water exposed to the sunshine, when a continuous series of small
bubbles of gas will at once be seen to escape from the intercellular
passages intersected by the cut. The gas thus evolved may be
collected with little trouble (Fig. 200), and will be found to be
chiefly
220 BOTANY PART i

oxygen, containing, however, under such simple conditions of experi-

ment, traces of other gases derived by diffusion from the plant and
the water. As water absorbs much less oxygen than carbonic acid (at
a temperature of 14 C. 100 vols. of water will dissolve only 3 vols.
of oxygen, but 100 vols. of carbonic acid), the escaping bubbles of

oxygen become visible; whereas the flow of the carbonic ;icid dis-
solved in the water to the assimilating plant is imperceptible.

Artificially conducting carbonic acid through the water increases, to a certain

degree, the evolution of oxygen, and thus the assiniilatory activity. Similarly an
artificial increase of carbonic acid in the air is followed by increased assimilation.

According to KREUSLER assimilation in sunshine attains its maximum in air con-

taining about 10 per cent of carbonic acid with a higher percentage it begins to
;

decrease. If the amount of carbonic acid gas be increased three hundred times

(from 0*03 per cent to 10 per cent in the atmosphere), the formation of starch is
only increased 4-8 times, while an increase of the C0 2 to six times the normal
proportion results, according to H. BROWN, in the formation of six times as much
starch. A longer exposure to an atmosphere rich in CO., does not have a favourable
influence on the health of the plant it takes on abnormal
;
modes of growth, loses
the power of flowering, and thus makes it evident that it is suited to live in
air with the ordinary proportion of C02 . This may be termed the "harmonic"
40
optimum ( ).

Like other vital activities of the plant assimilation is dependent upon the
temperature, and at lower grades of temperature is not so great as the available
light and carbon dioxide would permit. In bright, warm weather, on the other
hand, the small amount of C02 in the atmosphere becomes the limiting factor.
Carbon monoxide (CO) cannot be utilisedby green plants it cannot take the
;

place of the carbon dioxide, and is poisonous to plants, though less so than to
animals.
Under the same externalconditions, the assiniilatory activity of different plants
may vary from internal causes. In the same time and with an equal leaf surface,
one plant will form more, and another less carbohydrate. In this sense, it is
customary to speak of a "specific energy of assimilation," which is partly due to
the different number and size of the chloroplasts, as well as to a difference in the
structure of the leaves, but, without doubt, has also its cause in the greater or less
intensity of the assimilatory process itself.

As examples of medium assimilatory activity, the leaves of the

Sunflower and Pumpkin may be cited. Under conditions favourable
for assimilation, the leaves of these plants, according to SACHS, form in
a summer day of fifteen hours about 25 grammes starch per square
metre, while Cafalpa, according to BROWN and ESCOMBE, forms about
3 grammes per hour and square metre of surface. (The carbon for
the formation of the starch was supplied in the first cited example
from 50 cubic metres of air. A room of 120 cubic metres would
accordingly contain enough carbonic acid for 60 grammes of starch.)
From these figures a faint conception may be gained of the enormous
activity of the assimilatory processes, of the vegetation of the surface
of the earth. The German wheat harvest for 1900 represented about
23,000 million kilos of assimilated substance.
SECT, ii PHYSIOLOGY 221

The Mechanism of Gaseous Exchange

The gaseous exchange between one of the lower plants (or a submerged Phan-
erogam) and the surrounding medium is carried on by diffusion through the whole
surface of the plant. In a more highly organised plant the exchange takes place
by means of the stomata. The escape of aqueous vapour and the entrance and
escape of carbonic acid gas and oxygen in the processes of assimilation and respira-
tion (p. 244) alike take place almost entirely through these openings. But in
spite of their enormous number, the total area of the stomatal apertures is only
about 1 per cent or little more of the whole surface area, for each individual
opening is very small (about O'OOOl sq. mm. in Helianthus). Bnowx and
ESCOMBE have, however, shown that the minuteness of the openings, combined
with the peculiar distribution of the stomata, results in a greatly accelerated rate
of diffusion. Diffusion through a number of fine apertures is much greater than
through a single aperture of the same total area. When the small openings are
placed about ten times their diameter apart the diffusion is nearly as rapid as when
no separating wall is present. The distribution of the stomata in the epidermis
very nearly meets these requirements. For example, a square metre of the surface
of a Catalpa leaf absorbs about two-thirds the amount of carbonic acid gas taken

up in an equal time by the same area of potash solution freely exposed to the air ( ).
41

The Utilisation of the Products of Assimilation

The Formation of Albuminous Substances. The chlorophyll

bodies supply plants with organic nourishment in the form of a
carbohydrate. Although the greater part of the organic plant sub-
stance consists only of carbohydrates, as, for example, the whole
framework of cell walls, yet the living, and consequently the most
important component of the plant-body, the protoplasm, is composed
of albuminous substances. These albuminous substances have a com-
position altogether different from that of the carbohydrates. In
addition to carbon, oxygen, and hydrogen, they also contain nitrogen,
sulphur, and phosphorus, the nitrogen indeed in considerable pro-
portion (about 15-19 per cent). THERE TAKES PLACE ACCORDINGLY
WITHIN PLANTS A NEW FORMATION OF ALBUMINOUS SUBSTANCES.
There are certain indications that this formation is, in part, accom-

plished within the green cells of the leaves, but it must also be carried
on in tissues devoid of chlorophyll.
As little is known concerning the process of the synthesis of the albuminous
substances of plants as concerning the formation of the carbohydrates from the
carbonic acid and water. It has generally been supposed that they are formed
from the carbohydrates and mineral substances already mentioned, since these are
known to be transported to the region where the formation of protoplasm occurs,
and are there consumed. The carbohydrates utilised in this process seem to be
principally GLUCOSE (both grape-sugar, dextrose, C 6 Hi 2 6 + H 2 0, and fruit-sugar.
Isevulose, C B H 12 O 6 ) and MALTOSE (C^H^On x H 2 0) together with benzole derivatives ;

whatever may be the form of the original carbohydrate, whether starch, inulin,
cane-sugar, reserve-cellulose, or glycogen, glucose or maltose is always first formed
from it.
-2-1-1 1JOTANV PAKT I

Tlie mineral nitrates, sulphates, and phosphates t.-ike part in the process, chiefly
in the form of potassium and magnesium salts. Nitrogen anil sulphur are liberated
from the nitrates and sulphates, with decomposition of the acid radicals while of ;

the phosphates, the acid group is utilised in the formation of nuclein. Iron, which
is an essential plants whether or not they contain chlorophyll, appears to
for all

enter into the composition of the nut-rein. According to recent experiments

calcium salts appear to play a more immediate part in the p rote id synthesis than
was formerly supposed to be the case. Their importance, for most plants, is due to
their functioning as a medium for conveying the mineral acids, and for neutralising
or precipitating oxalic acid (C.,HoO 4 ). The oxalate of potassium, which is lirst
fiirmi-dfrom tin- potassium nitrate, reacts with tin- calcium salts present, witli the
formation of calcium oxalate, which is only slightly soluble, and crystallises out as
it accumulates. Wherever the formation of albumen or nuclein takes place, oxalic
acid is formed, the calcium salts of which may usually be found in adjacent cells,
in the form of aggregates of crystals, raphides, or crystal sand.
On account of their occurrence and behaviour in plants, the amides and hexa-
carbon bases are regarded as preliminary stages in the formation of albuminous
substances. Among the amides ASPAIIAGIX, CoH (NH 2 )(CONHo)(COOH), is note-
:J

worthy on account of its wide distribution. It is present in abundance in Gramineae

and Leguminosae (one litre of sap from Bean seedlings contains about 12- 15 grammes).
In the Cruciferae and Cucurbitaceae it is "replaced by glutamin, while in the
Coniferae a hexa-carbon base (arginin, C B Hi 4 N 4 2 appears to play the same role.
)

Any real knowledge of the mode of origin and transformation into more complicated
albuminous compounds of these and similar nitrogenous substances (c.ij. betain,
leucin, tyrosiu, and allantoin) is at present wanting.
The colloidal nature of many albuminous substances, which hinders their
osmotic diffusion, is of importance since it facilitates their recognition by appro-
priate reactions and their localisation in the protoplasm ( 42 ).

Transfer of the Products of Assimilation

AVhen
colloidal proteid substances are to be conveyed through the
tissues, as, for example, from seeds rich in proteicls into the seedlings,
they are by a hydrolytic process decomposed into soluble sub-
first
43
stances. According to SCHULZE ( ), albumoses and peptone are first
formed, and these are then broken up into amides and hexa-carbon
bases, and sometimes even into compounds of ammonia. They are in
this diffusible form transferred to places where, in combination with

carbohydrates and mineral acids, they are used anew along with
carbohydrates and salts in the formation of albumen.
In addition to the transfer of nitrogenous constructive material
through the parenchymatous tissues, the LONG-DISTANCE TRANSPORT
OF THE READY-FORMED ALBUMINOUS SUBSTANCES seems to take place
through the open sieve-tubes of the bast. It appears to be in the
sieve-tubes, which contain, during life, albuminous substances, starch
grains, drops of oil and leptomin, that the conduction of organic
substances is effected from the leaves. The increased thickening of
the cortical layers observed just above wounds made by ringing trees,
 PHYSIOLOGY 223

is due to the interruption and detention of a flow of nourishing sap

through the phloem towards the roots.

The transfer of the carbohydrates through unbroken cell walls to
the various points of consumption can- only be accomplished when they
are in solution. In case they are not already dissolved in the cell
sap, they must first be converted into soluble substances capable of
osmosis. This is of the highest importance for the transfer and utilisa-
tion of starch and reserve cellulose (hemicellulose). The former is
converted by the influence of DIASTASE into glucose or maltose.
Diastase belongs to those peculiarly acting substances termed
KKUMKNTS or ENZYMES, which possess the remarkable power of
decomposing or transforming certain organic compounds without
themselves becoming changed or consumed in the process. By virtue
of this property they are enabled to transform unlimited quantities of
certain substances if the resulting product is continuously removed.
According to their most important physiological properties, diastatic,
peptonising or proteolytic, and inverting enzymes are recognised, which
act on starch, albuminous substances, and sugars respectively. These
groups do not, however, exhaust the various ways in which these
substances, which play such an important part in the chemical changes
taking place in the organism, act. Thus there are trypsin-like
ferments which dissolve albumen in an alkaline medium, ferments
which dissolve membranes of cellulose, wood, or chitin, those that
decompose glucosides and oils, and others that convert urea into
ammonium carbonate (ureases). These and other enzymes have been
found in plants, and there is no doubt that such bodies play a part in
many imperfectly understood processes, especially in Fungi. A point
which the enzymes named above have in common is their hydrolytic
mode of action. They introduce the elements of water into the
substance acted on, starch (C 6 H 10 5 ), for example, being converted
into soluble glucose (C H 19 O e ).

BuCHNER discovered that the expressed sap of the yeast-plant,

even after being filtered, is able to transform grape-sugar into alcohol
and carbonic acid. He ascribed this property to a special enzyme
termed ZYMASE (alcoholase). The name of OXYDASES is
given to
certain substances which are regarded as acting by introducing oxygen.

Enzymes are for the most part colloidal albuminous substances which are
formed from the protoplasm, and exhibit a so-called catalytic mode of action.
They are easily rendered inactive by poisons or by too high a temperature.
Inorganic substances (catalysators such as finely divided indium or platinum)
exhibit a similar catalytic action to enzymes the power the latter possess of exciting
;

fermentation is thus not due to any special vital property. They can act when
removed from the organism, and may be precipitated from solution and again dis-
solved without loss of power. Besides ferments which hasten chemical reactions,
others which arrest the reaction are known (paralysators), others oppose and inhibit
44
the action of catalytic ferments (anti-ferments) ( ).
224 BOTANY

Other substances similar to diastase, and also capable of transforming starch

into dextrin, maltose or glucose, are widely distributed throughout the vegetable
kingdom, and are classed together as diastatic ferments. They are especially
abundant in starchy germinating seeds, as well as in tubers and bulbs, in leaves
and young shoots. They have also been found, strange to say, in organs where
there was no starch for them to act upon. The diastatic transformation and dis-
solution of the starch is accomplished in a peculiar manner. The starch grain
is not dissolved as a homogeneous crystal, uniformly from the surface imv.-ir IN
but becomes corroded by narrow canals, until it is finally completely disorganised,
and falls into small pieces (Fig. 201).

The complete transformation of the starch formed in the chloro-

phyll corpuscles during the day, takes place, as a rule, at night for in ;

the daytime the action of the

diastatic ferment is counter-
balanced by the formation of
s new starch. The glucose
which is thus produced in the
leaves passes out of the meso-
phyll cells into the elongated
cells of the vascular bundle-
sheaths and from them into
;
the phloem. The glucose
|* and maltose are transferred
through the leaf-stalks into
the stem. Thence they are
conveyed to the young shoots
Fi<:. -'01. Different stages of corrosion shown by tlir

starch grains of germinating Barley. and buds or carried down to

the roots
in short, they are
;

finally transported to places where they are required for the nutrition of
the plant. The glucose and maltose often become converted into other
carbohydrates, particularly into starch, during their passage from one
organ to another. Starch thus formed from other carbohydrates, and
not directly by assimilation, is often referred to as TRANSITORY STARCH,
and is iisually distinguishable by the smaller size of the grains.
For the purpose not of transport but of protection against cold starch i.s some-
times transformed into sugar (sweetening of potatoes) or oil (in the cortex and
sometimes the wood of trees) ( 45 ).

The Storage of Reserve Material

All the products of assimilation are not at once consumed. In

spite of this, however, assimilation is continued, and the surplus

products beyond the requirements of immediate consumption are re-

moved from the place of formation and accumulated as RESERVE
MATERIAL for future use. In our herbs, bushes, and trees, as the
yearly growth and consequent consumption cease towards the end
of each vegetative period, and as the assimilating organs have by
SECT, ii PHYSIOLOGY 225

that time attained their greatest expansion and efficiency, the surplus
of reserve material is
greatest at the close of the season, and is
stored in special RESERVOIRS OF RESERVE MATERIAL. The growth of
the shoots and leaves of the next season and similarly the growth
of seedling plants is dependent on such reserve materials. Reserve
materials will accordingly be found stored in different forms in the
cells of the embryo, or in the surrounding tissues of the seed, in

underground rhizomes, tubers, bulbs, and roots, or in the cortical

layers, the medullary rays, the wood parenchyma (especially the
thin-walled fibres), and the medulla of persistent stems. Conveyed
to these depositories of reserve material, the glucose and maltose are
again converted into other carbohydrates, usually starch. In other
cases the reserve carbohydrates take the form of cane-sugar (the

sugar-beet contains 5-8 per cent, and selected varieties 1 8 per cent, and
in some cases even 21 and 26 per cent), inulin (Compositae, Cam-
panulaceae) or reserve cellulose (e.g. vegetable ivory in the fruit of
Phytelephas). Still more remarkable is the transformation of carbo-

hydrates into fats and oils, occurring in the ripe and ripening seeds
of many plants, in fruits (Olive, oil-palm), and also in strictly vege-
tative tissues. In winter the starch in the wood of many trees also
becomes converted into but in the succeeding spring it is again
oil,

changed to starch. It at the opening of the buds, converted

is finally,

into glucose or maltose, and conveyed by the transpiration current

to the young shoots. Other receptacles of reserve material contain
scarcely any carbohydrate, but on the other hand there is much more
albuminous matter in the form of thick protoplasm, aleurone grains,
protein crystals, and fats (seeds of Ricinus). That in germination
similar tissues with protoplasm, nucleus, cell walls, etc., are formed
from these different materials, seems to indicate that all these
constructive materials are of almost equal value to the plants. This
is due to the fact that plants can apparently without difficulty
transform the carbohydrates, fats, or albuminous substances one into
the other, a result not yet accomplished by chemical processes.

Other Products of Metabolism

The chemicalactivity of the vegetable cell is by no means exhausted in the
production of the substances mentioned the increasing number of chemical com-
:

pounds found to be derived from the first product of assimilation is a matter of con-
tinual surprise. Of most of them neither the manner of their formation nor their
full importance in metabolism is understood. The conditions are not even fully
known which are necessary for the formation and functional activity of the OIUJAXK;
ACIDS (malic, tartaric, citric, etc.) which may in part be considered as products of
imperfect respiration while they are produced by some Fungi to acidify the medium
in which they live and render it less favourable for competing organisms. The
same holds for the equally widely-spread tannins. The function of the GLUCOSIDKS
is also imperfectly understood. These are compounds of sugars with a number of
different substances. They are soluble in water, and by the action of ferments or
Q
226 BOTANY PART i

dilute acids are broken up into glucose and other derivative products. It is con-
ceivable that the formation of glucosides (and tannins) serves to locally fix
substances which otherwise would readily diosmose. It might even be suggested
that the jtolysaccharides, starch and cellulose, might be regarded as glucosides of
the sugars themselves. In the Amygdalaceae (Bitter Almonds, Plum seeds)
AMYGDALIN is found which on fermentation yields hydrocyanic acid in addition to
benzaldehyde. The Lima or Java beans (P/iaseolus lunatus), which of late years
have been used for feeding cattle, also liberate from a glucoside hydrocyanic acid.
In the wild form this may be in such quantity (0'16 per cent) that fatal poisoning
has resulted. From a similar cause Sorghum, Glyceria, and Lotus may be more or less
poisonous. Glucosides appear in the Solanaceae as the poisonous SOLANIN, in the
Cruciferae (mustard seeds) as MYKONIC ACID, in the bark of the Horse-chestnut as
extremely fluorescent .&SCULIN, in species of Digitalis as the poisonous DIGITALIS',
and in the cortex of Willows as SALICIN. Certain plants (Indigofera, Polygonum
tinctorium) contain INDICAN, the glucoside of indoxyl the latter substance is con-
;

verted by oxidation into indigo. Woad (Isatis tinctoria) contains, on the other
46
hand, the related glucoside ISATIN ( ). CONIFERIN, which is present in the cambial
sap of the Conifers, has recently acquired an economic value, as from it VANILLIN,
the aromatic principle of vanilla, may be artificially produced. In this process the
coniferin is decomposed, through the action of a ferment or acid, into glucose and
coniferyl-alcohol, by the oxidation of which its aldehyde, vanillin, is formed.
It is as yet unknown what part in the metabolic processes of plants is performed

by the BITTER PRINCIPLES, such as the LUPULIN of Hops, ALOIN of Aloes, ABSYNTHIX
of Wormwood. There is the same uncertainty with regard to the functions of the
ALKALOIDS. Since most alkaloids, STRYCHNINE, BRUCINE, VERATRINE, CONIINE,
MUSCARINE, ATROPINE, QUININK, MORPHINE, CODEINE, ACONITINE, COLCHICINE,
NICOTINE, PILOCARPINE, COCAINE, together with CAFFEINE (theiue) and THEOBRO-
MINE which are closely related to uric acid, and many others are violent poisons,
their vegetable bases and repugnant bitter principles furnish a certain protection
to plants against destructive animals. This, however, does not preclude the
possibility that they may at the same time have an important physiological
significance. Thus, according to TREUB, hydrocyanic acid plays the same part
in the formation and transport of proteids in Pangium edule as the amides do in
other plants. It must, however, be noted that alkaloids are also poisonous to
the protoplasm of the plant ( 4? ).
The COLOURING MATTERS AND ETHEREAL OILS, although in actual weight present
make themselves particularly noticeable to the senses of
only in small quantities,
sight and smell. They probably represent only bye- and end-products of meta-
bolism ; and, with the exception of chlorophyll, take no further part in the vital
processes of plants, except in so far as they are beneficial to the general well-being
by enticing (e.g. flowers, fruits) or repelling (e.y. by warning colours) animals.
Their cecological significance is accordingly much better known than their physio-
logical function. Just as the ethereal oils are frequently found in special excretory
receptacles, the resins, gum-resins, and gum-mucilages, which are also excretion
products, are usually deposited in canals or glandular cavities, and are often mixed
with ethereal oils. Whether their formation in the particular instances is necessary
for the carrying out of the normal processes of metabolism is altogether uncertain.

They are, at any rate, useful to plants when wounded, serving as a protection
against evaporation and the attacks of parasites. On a square centimetre of the
surface of the splint-wood of the Pine, sixty to seventy resin canals open, and the
wood contains, according to MAYR, 22 kilos, of resin in every cubic metre.
 PHYSIOLOGY -227

The significance of the so-called india-rubber (CAOUTCHOUC) and GUTTA-PERCHA

in the latex in the economy of the plant is still less known. In addition to these
substances, there also occur in latex, resins, ethereal oils, alkaloids (in opium),
leptomin, starch grains, and other carbohydrates, oil-drops, and albuminous
substances. The presence of these substances valuable as constructive material, and

occasionally also of active enzymes (peptonising ferments are found in the milky
juice of Ficus Carica and Carica Papaya), in the latex, gave rise to the suggestion
that the latex cells and tubes function in the transport of the nutrient matter.
Our present knowledge of these often caustic and poisonous saps is limited to their
external utility in the economy of plant life. By their obnoxious properties they
defend plants from the attacks of enemies. Also, in the event of plants being
wounded, the latex is pressed out either by the surrounding turgescent tissue or by
the tension of the elastic walls of its own cells, and forms, as it quickly coagulates
in the air, an efficient covering for the wound. In other plants, especially in trees,
wound-gum serves the same purpose. (Pp. 134, 151.)

Special Processes of Nutrition

Parasites, Saprophytes, Symbionts, and Insectivorous Plants.

The acquisition of organic nutritive substances through the activity of
assimilating green cells is the most frequent, and is consequently con-
sidered the normal method of plant nutrition. Other modes of
nutrition are only possible at the cost of organic substances already
produced by the assimilatory activity of green plants. Some plants
forego all attempts to develop an adequate chlorophyll apparatus, and
by so doing become unable to provide themselves with nourishment
from the inorganic matter about them.
Great numbers of such colourless plants derive their nourishment
from the bodies of dead animals and plants. All organic matter at
one time or another falls into the power of such plants as are devoid
of chlorophyll it is chiefly due to their decomposing
; activity in the
performance of their nutritive processes that the whole surface of the
earth is not covered with a thick deposit of the animal and plant
remains of the past thousands of years. These peculiar plants are not
satisfied with the possession of the lifeless matter alone they even ;

seize upon living organisms, both animal and vegetable, in their search
for food.
It is chiefly the vast number of Bacteria and Fungi which nourish
themselves in this way as PARASITES (upon living organisms) or as
SAPROPHYTES (upon decaying remains of animals and plants and other
organic substances). But even some species of the most widely
separated families of the higher phanerogamic plants have also adopted
this method of obtaining food.
As a result of this modification of their manner of life, the organi-
sationand functions of these higher plants have undergone the most
remarkable transformation. From the corresponding changes in their
external appearance, it is evident how far-reaching is the influence
228 BOTANY PA in t

exercised by the chlorophyll. With the diminution or complete

disappearance chlorophyll, and consequent adoption of a
of the
dependent mode of life, the development of large leaf surfaces, so
especially fitted for the work of assimilation, is discontinued. The
leaves shrink to insignificant scales, for with the loss of their assimi-
latory activity the exposure of large surfaces to the light is no longer
essential for nutrition. For the same reason active transpiration
becomes unnecessary ; the xylem portion of the vascular bundle
remains weak, and secondary wood is
feebly developed. In contrast to
these processes of reduction resulting from a cessation of assimilation,
there is the newly developed power in the case of parasites to penetrate
other living organisms and to deprive them of their assimilated products.
In saprophytic plants, however, where the question is merely one of
absorbing nourishment from organic remains, the external adaptations
for taking up nourishment continue more like those for absorbing
the mineral salts from the soil, for it then depends only upon an
intimate contact with the decaying substances.

Cuscuta europaea (Fig. 202), a plant belonging to the family of the Convolvu-
laceae, may be cited as an example of a parasitic Phanerogam. Although, through
the possession of chlorophyll, it seems to some extent to resemble normally assimi-
lating plants, in reality the amount of chlorophyll present is so small that it is
evident that Cuscuta (Dodder) affords an example of a well-equipped parasite.
The embryonic Cuscuta plantlet, coiled up in the seeds, pushes up from the
ground in the spring, but even then it makes no use of its cotyledons as a source
of nourishment they always remain in an undeveloped condition (Fig. 202 at the
;

right). Nor does any underground root system develop from the young rootlet,
which soon dies off. The seedling becomes at once drawn out into a long
thin filament, the free end of which moves in wide circles, and so inevitably
discovers any plant, available as a host, that may be growing within its reach.
In case its search for a host plant is unsuccessful, the seedling is still able to creep
a short distance farther at the expense of the nourishing matter drawn from the
other extremity of the filament, which then dies off (t) as the growing extremity
lengthens. If the free end, in the course of its circular movements, conies

ultimately into contact with a suitable host plant, such as, for example,
the stem of a Nettle or a young Willow shoot (Fig. 202 in the centre), it twines
closely about it like a climbing plant. Papillose protuberances of the epidermis
are developed on that side of the parasitic stem in contact with the host plan.t,
and pierce the tissue of the host. If the conditions are favourable, these FIIK-
HAUSTORIA by special organs of absorption, the HAUSTOKIA (If).
are soon followed
These arise from the internal tissues of the parasite, and possess, in a marked
degree, the capability of penetrating to a considerable depth into the body of the
host plant by means of solvent ferments and the pressure resulting from their own
growth. They invade the tissues of the host, apparently without difficulty, and
fasten themselves closely upon its vascular bundles, while single hypha-like
filaments produced from the main part of the haustoria penetrate the soft
parenchyma and absorb nourishment from the cells. A direct connection is formed
between the xylem and phloem portions of the bundles of the host plant and the
conducting system of the parasite, for in the thin-walled tissue of the haustoria
 PHYSIOLOGY 229

there now develop both wood and sieve-tube elements, which connect the corre-
sponding elements of the host with; those of the parasitic stem (Fig. 202 at the left).
Like an actual lateral organ of the host plant, the parasite draws its transpiration
water from the xyleni, and its plastic nutrient matter from the phloem of its host.
The seeds of Orolanchc (Broom rape), another parasite, only germinate when in

Fio.- 202. O'."-"'" MtrtgKUO. On the viiiht. ^erniinatiiu; * ilin^s. In the middle, a plant of
Csotai parasitic on a Willow twij; : V reduced leaves : III. flower-clusters. On the left, cross-
section of the host-plantW, showing haustoria H
of the parasite CHS, penetrating the cortical
]>arenehyim anil in intimate contact with the xyleni r and the phloem of the vascular <

bundles; displaced cap

.*,
of sheathing sclerenchyina.

contact with the roots of the host plant its haustoria only penetrate the roots,
;

and^only its light yellow, reddish-brown or amethyst-coloured flower-shoot' appears

above the surface of the ground. Orobanchc, like Cuscuta, contains a small
amount of chlorophyll. Both are dreaded pests they inflict serious damage upon ;

cultivated plants, and are difficult to exterminate.

Many exotic parasitic plants, especially the Rafflesiaceae, have become so
completely transformed by their parasitic mode of life that they develop no
apparent vegetative body at all but grow altogether within their host plant,
;
230 BOTANY I'AIIT I

whence they send out at intervals tlu-ir extraordinary flowers. In the case of
Pilosti/les, a which lives on some shrubby Leguminosae, the whole
parasite
vegetative body is broken up into .filaments of cells which penetrate the host
plant like the mycelium of a fungus. The flowers alone become visible and
4S
protrude from the stems and leaf-stalks of the host plant (Fig. 203) ( ).

Flo. 203. Branch of a leguminous plant from the surface of which the flowers of a parasitic
Solir.s) art- protiudinu.
plant (Pilostyles Ulei, (From GOEBEL'S Organogm.plni.)

In contrast to these parasites, which have come to be absolutely

dependent upon other plants for their nourishment, there are certain
parasites which, to judge by external appearances, seem to be quite
independent, for they possess large green leaves with which they are
able to assimilate vigorous!}'. In spite of this, however, the plants
only develop normally, when their root system is in connection with
the roots of other plants by means of disc-shaped haustoria. They
may even (as is also the case with Cuscuta) enter into this relation
 PHYSIOLOGY 231

with other individuals of the same species. Thesium, belonging to the

Santalaceae, and the following genera of the Ebinanthaceae, Rhinan-
thits, Euphrasia, Pedicularis, Bartsia, and Tozzia may be mentioned as

examples of plants showing these peculiar conditions. In Tozzia the

parasitism is well marked in the earliest developmental stages. The
Mistletoe (Viscum album), although strictly parasitic, possesses, never-
theless, like many of the allied foreign genera of the Loranthaceae,
fairly large leaves well supplied with chloro-
phyll, and fully able to provide all the
carbohydrates required. It obtains, how-
ever, from the host plant (as HEINRICHER
has also shown to be probable in the case
of the Ehinanthaceae) its supply of water
and dissolved salts. Another member of the
Rhinanthaceae, Melampyrum, though prim-
arily a root parasite, has, on the other hand,
49
adopted a saprophytic mode of life ( ).
Humus plants, like some of the Orchi-
daceae (Neottia, Coralliorrhiza, etc.), and the
Monotropeae, are restricted to a purely
saprophytic mode of nutrition, and to that
end utilise the leaf -mould accumulated
under trees.
The roots and rhizomes of these saprophytes stand
in most intimate relation with fungal hyplise. Tlie
same indeed holds for the majority of green plants
which grow in woods and heaths, where the soil is
rich in humus. The fungal hyphse are sometimes
present in coiled masses within the cells of a definite
zone of the cortex, only occasional filaments passing
outwards to the soil. In other cases the fungus
surrounds the young roots with a dense investment
of interwoven hyphae. The former arrangement i*
spoken of as E~NDOTROPHIC, the latter as EXOTROPHIC
MYCOKHIZA the two types are, however, connected
;
FIG. 204. A root of Vicia Fvba,
by intermediate forms. A direct exchange of sub- with numerous root -tubercles.
stance between the soil and the root would appeal-
(Reduced.)
impossible in the case of the exotrophic mycorhiza.
On this ground, and because, despite the reduced surface exposed to the humus-
containing soil by the roots or rhizomes of total saprophytes, these obtain sufficient
nutriment, a co-operation of the fungus in the nutrition of such plants has
been assumed. The results of culture experiments lend further
support to this
view. Little is known, however, as to the nature of the relation between the
fungus
and the saprophyte. The view of PERCY GROOM and JANSE that in the case of endo-
trophic jnycorhizas the fungus supplies combined nitrogen, i.e. is ultimately digested
by thejoot, is supported by the work of W. MAGNUS, SHIBATA, and HILTNER. STAHL
on the other hand sees its use in a better
supply of salts from the soil. The latter
view, however, could only be a sufficient explanation in the case of green plants ( so ).
232 BOTANY PART I

The relation betweeu roots and certain Bacteria in the case of the Leguminosae is
better understood. It has long been known that peculiar outgrowths, the so-called

ROOT-TUBERCLES, are found on the roots of many Leguminosae (Bean, Pea, Lii]>in<',
Clover, etc.) (Fig. 204). These tubercles, of which a single bean plant may bear
4000, are caused by certain Bacteria (Bacillus radicicola, the two forms of which
are distinguished by HILTXER as Rhizobium radicicola and R. Beyerincki). These
Bacteria penetrate through the root - hairs into the cortex of the roots, and
there give rise to the tubercular growths. These tubercles become filled with
a bacterial mass, consisting principally of swollen and abnormally developed
(hypertrophied) BACTEUIOIDS, but in part also of Bacteria, which have remained
in their normal condition. The former seem to be eventually consumed by the
host plant, while the latter remain with the dead roots in the soil, to provide for
future reproduction. As the experiments of HBLLRIEGEL and the investigations of
NOBBE, BEYERINCK, HILTNER, and others prove, we have here a case of mutual
parasitism like those termed symbiosis by DE BARV. While the Bacterium lives
on carbohydrates and at first also on albuminous substances supplied by the host
plant, the latter profits by the power of fixing free nitrogen possessed by the
Bacteria. This is effected by means of an enzyme demonstrated by HILTNER. "While
the Bacteria remain alive they furnish a steady supply of nitrogenous substance to
the leguminous plant, and ultimately the remaining substance of the degenerated
Bacterioids is absorbed. When the large amount expended on nitrogenous
manures (Chili-saltpetre and ammonium sulphate) is borne in mind the agricultural
importance of this natural fixation of nitrogen will be evident. It has been

attempted to further it by infecting fields with soil rich in the bacteria or with
pure cultures of specially active forms (" nitragin"). The fact that the tubercle-
forming Leguminosae, unlike most plants, could flourish on ground poor in nitrogen,
and could accumulate stores of reserve proteids was known in the time of Pliny,
and these plants have long been known to form crops which enrich the soil by this
accumulation of nitrogen ( M ).
If the soil in which such a leguminous plant grows contains a sufficiency of

nitrates, few tubercles are formed on the roots. A similar immunity against infec-
tion is obtained by plants which bear a number of actively functional tubercles.
In addition to the Leguminosae (in which order only Gleditschia triacanthos has as
yet been found free from tubercles) tubercle formation due to a symbiosis with a
lower organism is known in Elaeagnus and Alnus. According to NOBBE and
HILTXER, these plants can also utilise free nitrogen. These authors have also
shown experimentally that the same holds for Podocarpus, which possesses
mycorhiza. Nitrogen-fixing Bacteria such as Azotobactcr chroococcum are also found
according to BENECKEand KEUTNEU in the sea according to REIXKE they lodge in
;

numbers in the mucilaginous coatings of Sea-weeds.

While among the higher plants only isolated forms have become
total or saprophytes, while
parasites in others the parasitism or
saprophytism is occasional or partial, among the lower plants large
families with numerous
genera and species are found completely
devoid of chlorophyll (Fungi and Bacteria), and wholly parasitic
or saprophytic in their mode of life. Of the Fungi and Bacteria
some are true parasites, and are often restricted to certain special
or animals, or even to distinct
plants organs ; others, again* are
strictly saprophytic in their habit, while others may be either
 PHYSIOLOGY 233

parasitic or saprophytic, according to circumstances. What renders

the conduct of these lower organisms particularly striking is the
peculiarity possessed by many of them of not fully utilising all of
the organic matter at their disposal but, on the contrary, so
;

decomposing and disorganising the greater part of it by. their

fermentative activity that, while at first competitors are inhibited,
later their own development soon becomes restricted. When a
mould attacks an apple, it not only takes the small amount of
organic matter necessary for its sustenance, but at the same time
converts the whole apple into a soft decaying mass upon which the
fungus itself cannot exist. A
considerable degree of heat is also
evolved in the course of these processes. The utilisation of the heat
produced in decomposing manure in making hot-beds is a familiar
practice. The heat produced by damp fermenting hay or raw cotton
may often become so great that in presence of inflammable gases
spontaneous combustion ensues. In germinating Barley an increase
in temperature of from 40 to 70 or more degrees has been observed.
The development of so much heat in this case is not due solely to
the respiration of the barley seeds, but according to CORN, to the
decomposing activity of a fungus (Aspergillus fumigatus). The
spontaneous combustion of raw cotton is, on the other hand, caused
by a Micrococcus. Coagulated albumen and thick gelatine are
rendered fluid by many Fungi and Bacteria, while the escaping gases
(carbonic acid, sulphuretted hydrogen, ammonium sulphide, ammonia,
etc.)show how far-reaching the decomposition is. In the same way
Penicillium brevicaule and other Fungi can liberate poisonous arsenical

gases when
living on a substratum containing arsenic such as the
green pigments of wall-papers. In this way by Gosio's method
minute quantities of arsenic can be detected ( v2 ). It is by similar
processes of decomposition that dead organic matter becomes
thoroughly disorganised and rendered harmless. To the poisonous
bye-products formed by Fungi and Bacteria is due the severity of
many diseases which they produce in living organisms (potato
disease, wheat smut, cholera, typhus, diphtheria, anthrax, etc.).
By the possession or formation of substances, which react as specific
poisons upon the infecting Bacteria, plants, and particularly animals,
in turn protect themselves against the attacks of such micro-organisms.
It is due to a knowledge of this fact that the science of Therapeutics
has been enabled to cope more and more successfully with infectious
diseases.

Fungi and Bacteria, in addition to the power, dangerous to themselves, of dis-

organising their own nutrient substratum by fermentation and putrefaction, also
possess the capability of making an unsuitable substratum suitable for their sus-
tenance. By means of inverting ferments they can convert an unsuitable cane-sugar
into an available grape-sugar, and by their diastatic ferments they are able to form
glucose and maltose from starch, and even from cellulose.
234 BOTANY PAKT i

Many Fungi and Bacteria are practically omnivorous, while others are extremely
exclusive in their selection of a substratum. Thus from ordinary tartaric acid
Penicillium only utilises the dextro-rotatory form, and Bacillus subtilis only llie
laevo-rotatory form. Aspergillus growing in a mixture of glucose and glycerine
"
utilises the former first ("election of nutritive materials). If the glycerine alone
is given completely utilised.
it is

As evident from their thriving upon such various substrata, Fungi have the
is

power of producing from the different carbon compounds (and also from nitrogenous
mineral compounds, such as ammonium tartrate, or even ammonium carbonate)
protoplasm, cell wall, nuclein, fat, glycogen, etc.

While many Fungi greater injury upon their host plants

inflict far

.by the decomposition they induce than by the withdrawal of the

nutritive substances, others produce a different effect. The Rust-fungi,
and the Agaricinae which attack trees, for instance, do comparatively
little injury to their host ; while the relation between host and fungus
in the case of the Lichens has been shown to be absolutely beneficial.
The Lichens were formerly considered to be a third group of the
lower Cryptogams and of equal value with the Algae and Fungi.
It is only comparatively recently that the discovery was made by DE
BARY and by the investigations particularly of SCHWEXDENER
verified
and STAHL, that the body of the Lichens is not a single organism,
but in reality consists of Algae (e.g. Cyanophyceae), which also exist
in a free state, and of Fungi, which for the most part belong to the

Ascomycetes these Fungi only exceptionally exist apart from the

;

Lichen. The fungal hyphre within the Lichen weave themselves

around the Algcxe and while the latter occupy the upper or outer
;

side of the leaf-like or cylindrical thallus as the more favourable

position for assimilation, the hyphae come into the closest contact with
them and absorb from them part of their assimilated products. The
fungi in return provide the Algae with water, which, owing to the
acids excreted, is often rich in nutrient salts, and enable them to live
in situations in which they could not otherwise exist. The researches
of ARTARIS make it probable that peptone is also provided by the

fungus. As a result of this close union with the fungi, the Algae
are in no way exhausted, but become more vigorous than in their free
condition, and reproduce themselves by cell division. As both sym-
bionts, the Algae as well as the Fungi, thus derive mutual advantage
from their consortism, Lichens form one of the most typical examples
of vegetable symbiosis ( 53 ).

The significance of the Cyanophyceae Nostoc and Awilmfmt in the roots of the
M
Cycadaceae and in the leaves of Azolla and other water plants is still unknown ( ).
In connection with these cases of symbiosis between plants, mention may here
be made of the similar symbiotic relation existing between animals and plants.
Like the Lichen-fungi, lower animals, according to BRANDT, profit by an association
with unicellular Algae by appropriating their assimilated products without at the
same time disturbing the performance of their functions. Freshwater Polyps
(Hydra), Sponges (Spongilla), Ciliata (Stetitor, Paramecium], also Heliozoa, Planaria,
 PHYSIOLOGY 235

and Amoebae (A. proteus) are often characterised by a deep green colour, due to the
numerous Algae which they harbour within their bodies, and from the products of
whose assimilation they also derive nourishment. In the case of the Radiolariaus,
the so-called "yellow cells," which have been distinguished as yellow unicellular
Seaweeds, function in the same way as the green Algae in the other instances.
Another remarkable example of symbiosis in which the relationship is not one
merely of simple nutrition, has been developed between certain plants and ants.
The so-called AST-PLANTS (Myrmeeophytea) offer to certain small, extremely
warlike ants a dwelling in convenient cavities of the stems (Cecropia), in branches
(Triplaris) in hollow thorns (Acacia spadicigera and sphaerocephala, Fig. 205), in
swollen and inflated internodes (ffumboldtia laitrifolia, Fig. 206), or in the
labyrinthine passages of their large stem-tubers (Myrmecodia, Fig. 207). At the
same time the ants are provided with food in the case of the Cecropias and Acacias

Jl
FIG. 205. Acacia sphaerocephaht. I, Leaf and part of stem S, hollow thorns in which the ants
;

live; F, food bodies at the apices of the lower pinnules N, nectary on the petiole. (Reduced.)
;

//, Single pinnule with food-body, F. (Somewhat enlarged.)

in the form of albuminous fatty bodies ("food bodies," Fig. 205 F), and by the
Acacias also with nectar (Fig. 205, A"). The ants in exchange guard the plants
most effectively against the inroads of animal foes as well 'as against other leaf-
cutting species of ants, which, in the American tropics, kill trees by completely
and rapidly divesting them of their entire foliage. These leaf-cutting ants, as was
shown by BELT and MOLLER, live in symbiosis with a Fungus (Rozites gongylophora).
Upon the accumulated leaves ("Fungus-gardens"), the ants make pure cultures
of a fungus mycelium, whose peculiar nutritive outgrowths serve them exclusively
for nourishment. Termites have more recently been discovered to be Fungus culti-
55
vators ( Other familiar examples of symbiosis are those existing between flowers
).

and birds or insects. The flowers in these instances provide the nourishment,
usually nectar or pollen, but sometimes also the ovules (Yucca-moth and the gall-
wasp of the Fig), while the animals are instrumental in the pollination. Here
also eachsymbiont is dependent upon the other. In the case of the unintentional
dissemination of fruits and seeds by the agency of animals, the symbiotic relations
are less close.

Of all the different processes of supplementary nutrition employed

230 BOTANY PART 1

by plants, those exhibited by Carnivorous or Insectivorous Plants in

the capture and digestion of animal.-; are unquestionably the most
curious. Although they are green plants and able to manufacture
carbohydrates for themselves they have, in addition, secured for
themselves, by peculiar con-
trivances, an extraordinary
source of nitrogenous organic
matter, by means of which
they are enabled to sustain a
more vigorous growth and
especially to produce more
seeds than would otherwise
be possible without animal
nourishment.

It is not accidental that the plants

which have become carnivorous are,
for the most part, either inhabitants
ofdamp places, of water or swamp*.
and moist tropical woods, or that
they are epiphytes. The nitrogenous
and phosphoric salts of the soil are
not obtained by them in the same
quantities as in the case of the more
vigorously transpiring land plants.
This is very evidently the case iu
the Sundew
(Droscra), which is
loosely attached by a few roots upon
a thick spongy carpet of Bog-nio-s
and must find in the animal food a
valuable addition to its nitrogenous
nourishment.
A great variety of contrivances
for the capture of insects are made
use of by carnivorous plants. The
leaves of Drosera are covered with
stalk-like outgrowths ("tentacles "),
the glandular extremities of which
Fio. 200. tfummxata iourUwta. showing the entrance ,. , -j -j A-
<l" cl ar g e a VISCld acld secretion
the hollo* tetemodT (From Scrap**
'
to
I'l.mt-Geography.) (Fig. 208). Any small insect, or even
larger fly or moth, which comes in
contact with any of the tentacles is caught in the sticky secretion, and in its in-
effectual struggle to free itself it only comes in contact with other glands and is
even more securely held. Excited by the contact stimulus, all the other tentacles
curve over and close upon the captured insect, while the leaf-lamina itself becomes
concave and surrounds the small prisoner more closely. The secretion is then dis-
charged more abundantly, and contains, in addition to an increased quantity of
acid, a peptonising ferment. The imprisoned insect, becoming thus completely
covered witli the secretion, perishes. It is then slowly digested, and, together
with the secretion itself, is absorbed by the cells of the leaf.
SECT. II PHYSIOLOGY 237

\\iPinguicula it is the leaf-margins which fold over any small insects that may
be held by the minute epidermal glands. In species of Utricularia (Fig. 47),
growing frequently in stagnant water,small green bladders (metamorphosed leaf-seg-
ments) are found on the dissected leaves. In .each bladder there is a small opening
closed by an elastic valve which only opens inwards. Small snails and crustaceans
can readily pass through this opening, guided to it by special outgrowths but ;

their egress is prevented by the trap-like action of the valve, so that in one

FIG. 207. Myrmecoitia n-hhu'tn. Tulirnms ^ti-m cut through longitudinally. Another epiphyte
is seen below. (J nat. size
; from .SCHIMI-ER'S Plant-Geography.)

bladder as many as ten or twelve crustaceans will often be found imprisoned at

the same time. The absorption of the disorganised animal remains seems to be
performed by forked hairs which spring from the walls of the bladder.
More remarkable still, and even better adapted for its purpose, is the mechan-
ism exhibited by some exotic insectivorous plants. In the case of Venus' Fly-trap
(Dionaea], growing in the peat-bogs of North Carolina, the capture of insects is
effectedby the sudden closing together of the two halves of the leaves (Fig. 209).
This action is especially due to the irritability of three bristles on the upper side
of each half-leaf (the leaf surfaces themselves are much less sensitive). Upon the
238 BOTANY

death of the insect caught by the leaf, a copious excretion of digestive sap takes

FIG. 2ns. Leaves of Droxera rotttndifolia. That on the left is viewed from above, that DM the
right from the side. (At'tei DARWIN enlarged.)

Fio. 209. A leaf of Dionaca muscipula, showing the

sensitive bristles on its upper surface, which, in the
parts shaded, is also thickly beset with digestive
glands. (After DARWIN, enlarged.)

place from glandular hairs on the leaf surface,

followed by the absorption of the products of
the digestive solution. In the case of other
well-known insectivorous plants (Nepenthes,
Cepkalotus, Sarracenia, Darlingtonia), the traps
for the capture of animal food are formed by
the leaves which grow in the shape of pitchers
(Figs. 46, 210). These trap-like receptacles are F
partially filled with a watery fluid excreted from 1

Fi<;. L'ln. 1'itrheivd leal of a tfeptn&O,

glands on their inner surfaces. Enticed by
A portion of the lateral wall of the
secretions ofhoney to the rim of the pitcher pitcher has been removed in order to
(in the case of Nepenthes),and then slipping show the fluid (f). excreted liy the
on the extraordinarily smooth surface below leaf-glands. (Red need.)
 PHYSIOLOGY 239

the margin, or guided by the downwardly -directed hairs, insects and other small
animals finally fall into the fluid and are there digested by the action of
ferments and acids. The larvae which CLAUTRIAU found living in Nepenthes
pitchers may, like intestinal parasites, be enabled to live under the conditions by
secreting a protective anti-ferment. In Sarrcfcenia and Cephalotus, GOEBEL was
unable to discover any digestive ferments but in Cephalotus it was possible to
;

determine that the secretions have antiseptic properties. The lid-like appendage
at the opening of the pitcher of Nepenthes, Sarracenia, and Cephalotus does not
shut ;
its function seems to be merely to prevent foreign substances from

('ailing into the pitcher, and particularly to keep out the

rain. The entrance to
the tubular leaves of Durlinylonia is under the helmet-like extremity, and there-
x
fore a lid is unnecessary ( ).

III. Respiration

It is a matter of common knowledge that animals are unable to

exist without breathing. In the higher animals the process of respira-
tion is so evident as not easily to escape notice, but the fact that
plants breathe is not at once so apparent. Just as the method of the
nutrition of green plants was only discovered by experiment, so it
also required carefully conducted experimental investigation to demon-
strate that PLANTS ALSO MUST BREATHE IN ORDER TO LIVE ; that, like
animals, they take up oxygen and- give off carbonic acid. Although
the question had already been thoroughly investigated by SAUSSURE
in 1822, and by DuTROCHET in 1837, and its essential features

correctly interpreted, LIEBIG pronounced the belief in the respiration

of plants to be opposed to all facts, on the" ground that it was
positively proved that plants on the contrary decomposed carbonic
acid and gave off the oxygen. He asserted that it was an absurdity
to suppose that both processes were carried on at the same time; and
yet that is what occurs.
ASSIMILATION AND RESPIRATION ARE TWO DISTINCT VITAL PRO-
CESSES CARRIED ON INDEPENDENTLY BY PLANTS. WHILE IN THE
PROCESS OF ASSIMILATION GREEN PLANTS ALONE, AND ONLY IN THE
LIGHT, DECOMPOSE CARBONIC ACID AND GIVE OFF OXYGEN, ALL PLANT
ORGANS WITHOUT EXCEPTION BOTH BY DAY AND BY NIGHT TAKE UP
OXYGEN AND GIVE OFF CARBONIC ACID. Organic substance, obtained
by assimilation, is in turn lost by respiration. A seedling grown in
the dark so that assimilation is
impossible, loses by respiration a con-
siderable part of organic substance, and its dry weight is consider-
its

ably diminished. It has been found that during the germination of

a grain of Indian Corn, a full half of the organic reserve material is
consumed in three weeks. That green plants growing in the light
accumulate a considerable surplus of organic substance is due to the
fact that the daily production of material by the assimilatory activity
of the green portions is greater than the constant loss which is caused
240 BOTANY PART i

by the respiration of all the organs. Thus, according to Boussix-

GAULT'S estimates, in the course of one hour's assimilation a plant
of Sweet Baywill produce material sufficient to cover thirty hours'

respiration. Plants produce in twenty-four hours about five to ten

times their own volume of carbonic acid. In shade-plants, according
to GRIFFON, this is usually reduced to twice the plant's volume,
while the commonly cultivated Aspidistra produces only one-half
its own volume, and can therefore succeed even under conditions
"
>T
which are unfavourable to assimilation (
:

).

A
means of judging of the importance of respiration is afforded by
the behaviour of the plants themselves when deprived of oxygen. By
placing them, for example, under a jar containing either pure nitrogen
or hydrogen, or in one from which the air has been exhausted, it will
be found that all vital activity soon comes to a standstill ; plants,
previously growing vigorously, cease their growth ; the streaming
motion of the protoplasm in the cells is suspended, as well as (with
few exceptions, e.g. Mimosa and Drosera) all external movement of the
organs. If oxygen be admitted, after not too long an interval, the

interrupted performance of the vital functions is again renewed.

A longer detention in an atmosphere devoid of oxygen will, however,
irrevocably destroy all traces of vitality ; as in every condition of
rigor internal chemical changes take place which, by a prolonged
exclusion of oxygen, lead to the destruction and disorganisation of
the living substance. THE PRESENCE OF OXYGEN is NECESSARY TO
THE CHEMICAL PROCESSES TAKING PLACE WITHIN THE CELL, IN ORDEU
TO MAINTAIN THE LIVING SUBSTANCE IN A CONDITION OF NORMAL
ACTIVITY.
The absorption of oxygen and the evolution of carbonic acid by living plants
can be demonstrated both qualitatively and quantitatively by simple experiments.
From what has already been said of the antagonistic nature of assimilation and
respiration, it will be at once apparent that these experiments must be conducted
either in the dark or on portions of plants devoid of chlorophyll. The more
abundant the protoplasm and the more energetic its vital activity, so much the
more vigorous is the respiration. The best results are obtained, therefore, from
young portions of plants in an active state of growth. It should also be mentioned
that in the following experiments only the carbonic acid and not the whole of
the products of the respiratory activity are determined. From theoretical con-
siderations, and also from exact chemical analysis, it has been definitely established
that, IN ADDITION TO CARBONIC ACID, WATER IS FORMED FROM THE OllUANH'
MATTER BY RESPIRATION.
The absorption of oxygen and the formation of carbonic acid may be clearly
shown by the following experiment (Fig. 211). A flask (J?) partly filled with young
mushrooms or flower heads of a Composite is inverted with its mouth in an open
vessel of mercury (Q), and a few centimetres of caustic potasli solution (AT) introduced
within its neck. In the same degree as the carbonic acid produced by respiration is
absorbed by the caustic potash, the volume of air in the flask will be reduced and
the mercury will rise in the neck. After a time, the ascent of the mercury ceases
and it remains stationary. If the quantity of air remaining in the flask be esti-
SF.CT. ii PHYSIOLOGY 241

mated, it will be found that it has lost a fifth of its original volume ;
this means
that the whole of the oxygen (which makes up one-fifth of the atmospheric air) has
been absorbed. If caustic potash is not used in this experiment to absorb the
exhaled carbonic acid, the mercury remains at its original level, or, in other words,
the volume of air in the flask remains unchanged. From this experiment it is
apparent that the volume of oxygen absorbed is equal to the volume of carbonic
COo
- 2
acid evolved, as expressed by the formula -

=l. This equivalence of volume

U.,
between the oxygen absorbed and the carbonic acid exhaled exists only in cases
where the oxygen is used exclusively for complete respiration, and not where it is
consumed in transforming the contents of the cells, as is observed in the germination
of seeds rich in fat, and in the interchange of gases in the case of the succulents. In
the germination of seeds rich in fat, the fat is converted into carbohydrates richer
in oxygen. The oxygen consumed remains combined in the plant. On the other
hand, in the case of the succulents the production of C0 2 falls with the formation
of organic acids (malic acid in the case of the Cactaceae, oxalic acid in the Mesem-

bryanthaceae) in the dark these acids are, however, again decomposed in the
;

light and, according to AD. MAYER, starch is formed and oxygen set free. The
respiratory co-eflicient may further vary within certain limits according to the
nutrition and conditions of cultivation of a plant.
The absorption of oxygen in the respiration of plants can also be shown by the
fact that a flame, held in a receptacle in which plants have been kept for a
time, is extinguished. If a lighted taper be thrust into a glass cylinder which
has been partially filled with flowers or mushrooms and then tightly covered
and allowed to remain it will be extinguished, as the oxygen of the
for a day,
air in the cylinder will all have been absorbed. The carbonic acid exhaled in
respiration can be quantitatively determined from the increase in the weight of the
caustic potash by which it has been absorbed, or by conducting the respired carbonic
acid gas through baryta water and estimating the precipitate of barium carbonate.

Intramolecular Respiration (
58
).
In 1871 AD. MAYER showed
that the yeast-plant could obtain the chemical energy necessary for its
life from changes within its organic substance and without absorbing

oxygen. In the middle of the 'seventies PFLUGER made the discovery

that frogs are not only able to live for some time in an atmosphere
devoid of oxygen, but even continue to exhale carbonic acid. From
similar investigations it was found that the higher plants also, when
deprived of oxygen, do not die at once, but can prolong their life for
a time and evolve carbonic acid. Under these circumstances it is
apparent that both elements, the carbon as well as the oxygen, must
be derived from the organic substance of the plants themselves the :

oxygen can only be obtained through some unusual process of decom-

position carried on within the plant. This form of respiration has
consequently been described as intramolecular (anaerobic) respiration.

The amount of carbonic acid produced in a given time by intramolecular

respiration is usually less (|-) than that given off in the same time during normal
respiration. There are plants, however (for instance, Vicia Faba), whose seedlings,
in an atmosphere of pure hydrogen, will exhale for hours as much carbonic acid as
in the ordinary air. During intramolecular respiration in aerobionts all growth
R
242 BOTANY

ceases and abnormal processes of decomposition take place, whereby, as for instance
in alcoholic fermentation, alcoholand other products are formed.
Intramolecular respiration commences as soon as the access of free oxygen to
the protoplasm is prevented, and continues until tin- latter is killed by the accumula-
tion of injurious products of decomposition. The sufficiently early readmittance
of oxygen may, however, permit the cell to resume its normal condition ,by re-
establishing ordinary respiration and
removing the accumulated products
of decomposition.
Some plants endure the absence
of freeoxygen badly and only for a
short time, others better and for a

longer period. Certain of the lower

plants (Bacteria, Fungi, Characeae)
can exist for a considerable time
without free oxygen, or are even able
under favourable conditions to find in
intramolecular respiration a complete
substitute for the ordinary oxygen-
respiration. This capability is so ex-
treme in some Bacteria that no trace
of free oxygen is necessary for their
existence, while some cannot live in
the presence of oxygen. To distin-

guish such specially adapted organ-

isms from those which depend for
their normal respiration on free

oxygen (aerobionts or aerobes) they

are termed anaerobionts or anaerobes.
The grades of independence of free
oxygen indicated above may be dis-
tinguished as temporary or faculta-
tive anaerobiosis and permanent or

obligate anaerobiosis respectively.

Plants which can store up substances
rich in oxygen (e.g. certain pigments)
and at the expense of these live for
Km. 211. Experiment to demonstrate respiration.
a time without an external supply
The inverted tlask (IS) is partially filled with

(lowers, which are held in plai-e by the plug of of oxygen may be distinguished as
cotton (W). Through the absorption of the car- pseudo-anaerobes.
bonic acid exhaled in respiration, by the solution
Respiration as a Source of Energy.
,,,' eaustie

nerk of the
potash
flash.
(^
the mercury^) rises in the _ he lnetabolic c , CQU .
stituting respiration is a necessary
concomitant of life maybe inferred from the cessation of vital manifestations when
respiration is interfered with, and the association of more active life with increased
demands on the respiratory process.
In respiration chemical changes occur leading to the setting free of energy. It
is hardly too much to say that it is the energy obtained by respiration which serves
to carry on and maintain the vital manifestations. Thus a specific vital energy is
obtained by means of respiration, which might be termed vital force were this term
not used in another sense sucli a supply of energy is not provided by other
;
 PHYSIOLOGY 243

sources of force in the plant (force of pressure in turgescence, vibrations of light

and heat rays, etc.).
The majority of plants sacrifice in physiological combustion a portion of their

organic substance (especially carbohydrates) in order to obtain this driving power.

The combustion is as a rule so complete that CO., and H.,0 are the resulting products ;

this not only obtains the maximum of energy but obviates the accumulation of
injurious products of respiration. In other cases, however, considerable amounts of
organic acids are formed. This modification, which is found in succulents living under
unfavourable conditions for gaseous exchange and consequently for assimilation,
avoids the loss of carbon attendant on the liberation of C0.2 into the air. (Cf. p. 241).
The energy liberated by the respiratory combustion of carbon-compounds is
traceable back to that stored in the form of potential energy of chemical combination
in carbohydrates, which were formed by the help of the sun's rays in assimilation
(cf. p. 216). not only liberated in the more or less complete combustion
Energy is

of carbon-compounds but may be obtained in other chemical processes. While

most plants consume organic substances in respiration, some lower plants, especially
Bacteria, depend on energy obtained from other chemical changes. Thus sulphur-
bacteria oxidise sulphuretted hydrogen to sulphur, and this in turn to sulphuric
acid ; others, according to NATHAXSOHN, oxidise thio-sulphates. The nitrite-bacteria
form nitrous acid from ammonia and amides, while nitrate-bacteria convert the
nitrous into nitric acid. It is doubtful whether the iron-bacteria gain a similar
benefit from the oxidation of compounds of iron. The energy obtained in such
processes may serve to replace the driving power of respiration, or, as has been
shown for the uitro-bacteria (p. 189), and is possibly the case for the sulphur-
bacteria also, may effect the synthesis of organic carbon-compounds. The working
power is thus devoted to one or other process of the life of the plant (").

Fermentation and Respiration (). In the section on special

modes of nutrition it was stated that when this is effected at the
expense of organic food material, fermentation is frequently set up
in the substratum. Since fermentation is a process of decomposition
associated with a liberation of energy stored in organic compounds,
it
may be serviceable in the same way as respiration. Its amount
may also be more or less influenced by the respiratory needs of the
organism.

Thus, when the yeast plant is living as an anaerobe, the greater part of the
organic substratum (about 98-99 per cent) is fermented. When growing as an
aerobe, with a full supply of oxygen, a larger amount of the substratum is available
for use in the processes of growth and multiplication.
Since, even in the latter case, a large part of the grape-sugar (according to
BUCHNER and RAPP about 85 per cent) is fermented, it may be concluded that the
process of fermentation, though under certain circumstances a source of energy to
the plant, is to some extent independent of the respiratory needs of the latter,
and may have a further significance. Since the products of fermentation are as
a rule far better endured by the organisms which have given rise to them than
by others, their production may have an oscological value in the struggle with
competing organisms. Oxygen-respiration is also to a certain extent independent

of the other vital manifestations. It is most active at a temperature a little below

that which causes the death of the organism, when all other activities are being
arrested by the heat.
244 BOTANY PART i

Heat produced by Respiration. Respiration is, chemically and

physically considered, a process of oxidation or combustion, and, like
them, is accompanied by an evolution of heat. That this evolution
of heat by plants is not perceptible is due to the fact that consider-
able quantities of heat are rendered latent by transpiration, so that
transpiring plants are usually cooler than their environment ; and
also to the fact that plants possess very large radiating surfaces in
proportion to their mass.
The spontaneous evolution of heat is easily shown experimentally, if transpira-
tion and the loss of heat by radiation are prevented and vigorously respiring plants
are selected. Germinating seeds (Peas), if examined in large quantities, show under
proper conditions a rise in temperature of 2 C. The greatest spontaneous evolu-
tion of heat manifested by plants has been observed in the inflorescence of the
Araceae, in which the temperature was increased by energetic respiration 10, 15,
and even 20 C. Also in the large flower of the Victoria reyia temperature
variations of 15 C. have been shown to be due to respiration. One gramme of the
spadix substance of an Aroid exhales, in one hour, up to 30 cubic centimetres CO2 ;

and half of the dry substance (the reserve sugar and starch) may be consumed in a
few hours as the result of such vigorous respiration. In the process of wound -

healing in plants a noticeable rise in temperature also occurs.

That other processes, in addition to respiration, co-operate in the production
of heat is apparent from the fact that the amount of heat evolved does not vary
proportionally to the carbonic acid exhaled.

The Movement of Gases in Respiration

In plants of simple construction the cells which are in direct

contact with the air or water can absorb the requisite oxygen
directly ; while cells in the midst of tissues are dependent upon the
oxygen which can diffuse through the surrounding cells. Such a
diffusion from cell to cell would not, however, be adequate, in the case
of the large cellular bodies of the higher plants, to provide the living
cells of the interior with a sufficient supply of oxygen. This is
accomplished by means of the air-spaces, which, as INTERCELLULAR
PASSAGES, penetrate the tissues in all directions and so bring to the
protoplasm of the inner cells the air entering through the STOMATA
and LENTICELS. The path of the respiratory gases is thus the
same as that followed by the gases in transpiration and assimilation
(pp. 205, 221).

The movement of the gases within the intercellular spaces is due partly to the
diffusion, induced by the constant interchange of gases caused by respiration,
assimilation, and transpiration, and partly to movements in mass arising chiefly
from modifications of the temperature, pressure and moisture of the surrounding
atmosphere, but also increased by the movement of the plants themselves, through
the action of the wind.
That the intercellular spaces were in direct communication with each other and
 PHYSIOLOGY 245

also with the outer atmosphere was rendered highly probable from anatomical
investigation, and has been positively demonstrated by physiological experiment.
It is, in fact, possible to show that air forced by moderate pressure into the inter-
cellular passages makes its escape through the stomata and lenticels and con- ;

versely, air which could enter only through the stomata and lenticels can be drawn
out of the intercellular passages. The method of conducting this experiment can
be seen from the adjoining
figure (Fig. 212).
Intercellular air-spaces are
extensively developed in water
and marsh plants, and occupy
the greater part of the floating
portions of the plant. The sub-
merged portions of water plants
unprovided with stomata thus
secure a special internal atmo-
sphere of their own, with which
their cells maintain an active
interchange of gases. This in-
ternal atmosphere is in turn
replenished by slow diffusion
with the gases of the surround-
ing medium. In marsh-plants,
which stand partly in the air,
the large intercellular spaces
form connecting canals through
which the atmospheric oxygen
without being completely used
up can reach the organs growing
deep in the swampy soil and
cut off from other supplies of
oxygen. In some cases the
need of a supply of oxygen
to such roots is met by special-
FIG. 212. Experiment to show the direct communication
ised roots (pneumatophores) of the external atmosphere with the internal tissues of
which project vertically from plants. The ^lass tube R, and the leaf }', are fitted air-
the muddy soil (Fig. 213). tight in the bottle ;upon withdrawal of the air in the
Phosphorescence. Under bottle by suction on the tube B, the external air pene-
trates the intercellular spaces of the leaf, through the
the same conditions as the
stomata, and escapes in the form of small air-bubbles
respiratory process a limited from the cut surface of the leaf-i>etiole. (From DKTMER'S
number of plants, particularly Pliysiol. I'rakt.)

Fungi and Bacteria, emit a

phosphorescent light. This phosphorescence at once disappears in an atmosphere
devoid of oxygen, only to reappear on the admission of free oxygen. On this
account the phosphorescent Bacteria, according to BEYERINCK, afford a delicate
test for the activity of assimilation. All the circumstances which facilitate
respiration intensify phosphorescence the converse of this is also true. According
;

to the results of investigations concerning the phosphorescence of animals, from

which that of plants does not probably differ in principle, 'the phosphorescence
is not directly dependent upon the respiratory processes.
The best-known phosphorescent plants are certain forms of Bacteria which
246 BOTANY I'AKT I

"
occur in the sea and the mycelium, formerly described as Rhizomorpha,"
of the Fungus Ayaricus mellcvx.
As further examples of spontane-
ously luminous Fungi may be
cited Agaricus olearius, found grow-
ing at the foot of olive trees in
South Europe, Polyporus sulphureus
and other less familiar Agarics (Ag.
igncus, noctilucens, Gardncri, Mycena
illuminans, etc.). The phosphor-
escence of decaying wood is also,
without doubt, due to the growth
of Fungi or Bacteria, but some-
times to an insect (Neanura) which
reacts in this way when disturbed.
On phosphorescent fish or meat
harmless phosphorescent Bacteria
(Microspira photogena, Pseudomonas
lucifera) occur.According to
MOLISCH Micrococcus
phosphoreus
usually occurs on meat which has
been moistened with a 3 per cent
solution of common salt and kept at
a low temperature. Of plants taking
part in the phosphorescence seen in
the sea, the most important are
Pyrocystis noctiluca belonging to the
Gymnodiniaceae and certain Peri-
dineae. Their phosphorescence,
according to observations on Ccr-
atium tripos made by REINKE, is
Fio. 213. Respiratory root of Ai-'tctnnia, one of the
Mangroves, (i nat. size from SCHIMPER'S Plant-
;
brought about by mechanical as well
Geography.)
as by thermal and chemical stimuli.}
The so-called phosphorescence of
the Moss Schistostega, of the Flagellate Chromophyton Kosnnoffii, and of some

Fio. 214. Phosphorescent cell of the protonema of Schistostegn. S' S', Path of a ray of light \vliicti
after passing through the chloroplasts acquires a .un'i'iit
inge and is totally rrtlrrtrd. (Cf. Fig. 172.)
 PHYSIOLOGY 247

Selaginellas and Ferns, has nothing in common with actual phosphorescence,

but is produced solely by the reflection of the daylight from peculiarly formed
cells (Fig. 214). The phosphorescence observed in some seaweeds results, on the
other hand, from the fluorescence and opalescence of certain of their albuminous
61
substances, or from the iridescence of their cuticular layers ( ).
The light occasionally emitted by some floral structures on sultry nights is, we
must assume, electrical and comparable to St. Elmo's fire.

IV. Growth
The size which plants may attain varies enormously. A Mushroom
seems immeasurably large in contrast with a Micmcoccus, but small if
compared with a lofty Californian Sequoia. A Bacillus of the size of
a Mushroom, or a Mould-Fungus of the height of a Sequoia, are,
with their given organisation, physiologically as inconceivable as a
Mushroom with the minuteness of a Micrococcus. The size of an
organism accordingly is an expression of its distinct individuality,
and stands in the closest relation to structure and conditions of life,
and in the individuals of a species only varies within certain narrow
limits.
However large a plant may be, and however numerous its cells,
it nevertheless began its existence as a single cell, microscopically
small and of the simplest structure. To attain its final size and
perfect development it must grow, that is, it must enlarge its body

and undergo differentiation. Even for the minute unicellular Bacteria

growth is essential, as they multiply chiefly by cell division. Each
daughter cell must grow and attain the dimensions of the parent cell,
or in a few generations the capacity for existence itself will be lost
through their continually decreasing size. It is in fact impossible
to conceive of a plant where development is not the result of
growth. If a growing Oak or Cedar be compared with the single

spherical egg-cell from which it has arisen, it is at once clear that by

the term growth we mean not only an increase in volume, but include
also a long series of various developmental stages, and external
and internal modifications. A mere increase in volume does not
necessarily imply growth, for no one would say that a dried and
shrivelled turnip grows when it swells in water. On the contrary,
active growth may be accompanied by a considerable loss of
substance, asis shown by the sprouting of potatoes kept in a dark

cellar water is lost through transpiration as well as organic

;

substance through respiration, and yet the new shoots show true
growth.
In the lower organisms growth is exhibited in its most simple
form. In an amoeba or a plasmodium growth is simply an increase
in their substance in a unicellular Alga or in a Fungus it means, in
;

addition to this, an enlargement of their cell walls. In the higher

248 BOTANY I-AKT i

plants the processes of growth are far more complicated and various,
so that, according to SACHS, three chief phases of growth can be
distinguished, which, however, are not sharply separated, but merge
imperceptibly one into the other :

1. The embryonic phase in which the rudiments of new organs

are formed.
2. The phase of elongation of the already formed embryonal
organs.
3. The phase of internal development and completion of the
tissues.

The Embryonal Development of the Organs

Plants, in contrast to the higher animals, continually develop new

organs. These arise either from tissues retained in their embryonic
condition, as at the growing point, or they have their origin in

regions which have already more or less completely attained their

definite form. The leaves and shoots spring directly from the
tissues of the growing point ; the first lateral roots, however, make
their appearance at some distance from the growing point, where
a perceptible differentiation of the tissues has already taken place.
Leafy shoots may also take their origin from old and fully
developed tissues, which again assume an embryonic character,
accompanied by an accumulation of protoplasm and renewed activity
in cell division. But as this only occurs in exceptional cases, shoots
which thus arise out of their regular order are termed ADVENTITIOUS.
The manner of the Formation of New Organs at the Growing
Point has already been described in the morphological portion of this
book. It is only necessary here to again call attention to the fact
that the young organs, with few exceptions, develop in acropetal
succession, so that the youngest is always nearest the apex. The
point from which new organs arise, and the number which develop,
are chiefly dependent upon inherited internal disposition. External
factors can, however, exert an effect in particular cases. The
influence of such factors as light, gravity, chemical and mechanical
stimuli, which at certain times in the later life of the tissues is of
extreme importance, has usually but little effect on the embryonal
development. Yet, on the other hand, the position of the first
division wall of the germinating spore of Marsilia is determined by
the action of gravity, and the direction of the first wall (as well as
of the preceding nuclear division) in the spore of Equisetum and the
ova of Cystoseira barbata, Pelvetia, Haliseris, and Ascophyllum among
the Fucaceae, and of Dictyota is determined by its relative position to
the light ( 62).
In Adventitious Formations, on the contrary, the influence of
external forces is often very evident, as, for example, in the formation
SECT, ii PHYSIOLOGY 249

of climbing-roots, which in the Ivy and other root-climbers are de-

veloped only on the shaded side of the stem. In the Alga Caulerpa
the new leaf-like organs arise only on the illuminated side of the parent
organ. It is, on the other hand, the force of gravity which excites
the formation of roots on the under side of underground rhizomes. It
is also due to gravity that the growing points of shoots are formed only

from the upper side of the tubers of Thladiantha dubia, or that new
twigs develop, for the most part, from the upper side of the obliquely
growing branches of trees. Contact stimuli, on the other hand,
determine the primary inception and point of development of the
haustoria of Cuscuta. The sexual organs of Fern prothallia are always
developed on the side away from the light that is, in normal con-
;

ditions on the under side, but in case of artificial illumination on the

upper side.

As a result of one-sided illumination and the stimulus of gravity, together with

the favouring influence of moisture, the rhizoids spring only from the under side
of the gemmae of Marchantia, so that eventually the two originally similar sides
assume an altogether different anatomical structure.
Many adventitious formations are the result of definite external causes ; as, for
example, the galls induced by the stings of insects and the deposits of animal eggs
and larvae (cf. p. 165).

The development of adventitious formations is especially induced

by MUTILATION of plants the same effect may result when the organs
;

or growing points though uninjured become functionless. NEW

FORMATIONS are in this manner produced at points from which they
would never have arisen on the uninjured plants. In the case of
Pelargoniums, Willows, and many other plants, it is possible to
induce the formation of roots wherever the shoots are cut. In other
plants, however, there seem to be certain preferred places, such as
the older nodes, from which, under the same circumstances, roots
develop. In like manner new shoots will appear in the place of
others that have been removed. IN THE DEVELOPMENT OF NEW
FORMATIONS ON A MUTILATED PLANT THOSE VERY ORGANS ARISE,
OF WHICH THE PLANT HAS BEEN DEPRIVED. Root-less shoots
develop first of all new roots. Roots and root-stocks deprived of
their shoots form first new shoots. In these processes there is
manifested an internal reciprocity in the formative growth of organs,
which has been termed the CORRELATION OF GROWTH.
Correlation of growth is often, also, very apparent in the normal development
of the organs of uninjured plants. It is due to this that scales of buds are developed
in their special form rather than as foliage leaves. For, as GOEBEL showed, it is
possible by artificial means, as, for example, by the timely removal of the leaves of
the parent shoot of Aesculus, Acer, Syringa, tyuercus, or in the case of Prunus
Padus, by cutting off the upper extremity of the shoots, to induce the formation of
normal foliage leaves in the place of the scales. The vigorous growth which
ensues in the fruit and in the fruit-coverings after fertilisation and development
250 BOTANY i AKT i

of the embryo in the ovule, affords another example of correlation for,

;
in case no
fertilisation of the egg-cell occurs, all those changes which produce a ripe fruit
from the flower do not take place and, instead, another correlative process is
;

manifested by which the now useless organs are discarded. Certain plants, especi-
ally those modified by cultivation, form an exception to this: in many varieties of
Banana, in the seedless Mandarin, and in the variety of raisins known as Sultana,
etc., although no seeds capable of germination are produced, the formation of a
fruit is nevertheless continued. Even in these instances it is essential for the
formation of fruit that there shall have occurred a previous pollination of the
stigma, or the fertilisation of the ovules, which, however, do not mature. According
to MULLER THUKGAU ( 63 ) the formation of seeds in grapes exerts an influence on the
form, quantity, and quality of the succulent portion of the berry. In some few
exceptional cases, however, as in the fruits of the Fig and Gherkin and the seeds
of Cycads, even this impetus to fruit formation is not necessary. The manner of
the formation of conducting tissues in plants, and also their anatomical develop-
ment, are regulated by correlation. From these few instances it may be seen how
the principle of con-elation affects the most various of the vital processes, even
under normal conditions, and how the harmonious development and function of
the single members of the plant body are controlled by it.
The polarity manifested by plants should also be considered as a special example
of the correlation existing between the different parts of the plant body. This
polarity is particularly apparent in steins and roots, and finds its expression in the
tendency of every small piece of a stem to develop new shoots from that end which
was nearer the stem apex, while the roots take their rise from the other end.
Pieces of roots in like manner send out roots from the end originally nearer their
apex, and shoots from the end towards the stem.
In accordance with this principle, detached pieces of stems produce new shoots
from their " shoot-pole," and injured roots new roots from their " root-pole." This
polarity, particularly investigated by VOCHTING and SACHS, is supplemented accord-
ing to GOEBEI. by the nutritive current setting towards the wounded surface ;
it

makes apparent in even the smallest pieces of stems or roots, and may, in this
itself

respect, be compared to the magnetic polarity exhibited by every small piece of a

magnet. Unlike poles of a plant may readily be induced to grow together, while
like poles may only be brought to do so with difficulty, and then do not develop

vigorously. As a result of such experiments, a radial polarity has also been recog-
nised by Yik'HTlNu in stem and root tissue thus, for instance, pieces of a stem or
:

root, inserted in a lateral incision of a similar organ, become united with it, if they
are so placed that the side originally outermost occupies the same relative position
in the new organ, but if this position is altered no such union takes place. Leaves
take, in respect to polarity, a special position, for they are not organically included
within new formations derived from them. Thus, from the basal end of a leaf, an
entire plant, with roots, stem, and leaves, may arise, while the regenerative leaf
itself gradually dies. It is of especial interest to observe the effect of external
influences upon the position of new formations, when they come into opposition to
the internal disposition of the plants themselves. In this respect, the behaviour
of different species varies greatly. In one, the internal factors predominate, that
is, the new formations appear quite independently of external conditions
in ;

another, the external influences for the moment prevail, but the internal dis-
position of the plant, when thus constrained for the time being, ultimately makes
itself apparent and the new formations never develop vigorously. A willow twig,
planted in a reversed position, with the shoot-pole in the ground, will produce
 PHYSIOLOGY 251

roots, and from the root-pole may even produce shoots. These, however, usually
soon die and their place is supplied by other stronger shoots arising from the
shoot axis just above the roots. It is only by the most careful suppression of any
such developments that the shoots from the root-poles may be kept alive. In the
trailing shoots of such plants as the Blackberry roots may be formed beneath the
uninjured growing point. In so-called "weeping" trees, the formation of side
branches from the upper side of the hanging branches is favoured by external con-
but the internal polarity prevents their vigorous development, and those
ditions,
formed soon die. In the cultivation of vines and fruit trees this peculiarity is
utilised to produce short-lived, fruit-producing shoots by bending over the vines
or training the branches of the trees in the cultivation of wall fruits. On the
other hand, in some cases the internal polarity is easily overcome by external in-
fluences. It is sometimes sufficient merely to reverse the erect thallus of Bryopsis,
one of the Siphoneae, to convert the former apical portion into a root-like tube
which penetrates the substratum and fastens itself to the grains of sand. It has
also been positively determined, although otherwise such cases are unknown

among the higher plants, that the growing points of the roots of Neottia and of
certain Ferns (Platycerium, Asplenium esculentum) may be converted through some
inherent tendency into the vegetative cone of a stem (cf. p. 47) ( 84 ).
The correlation phenomena manifested in the formation of new organs have the
greatest practical importance, for the propagation of plants by cuttings or grafting
isbased upon them.
In artificial reproduction detached pieces of plants are made use of for the
purpose of producing a fresh complete plant. In many cases this is easily done,
but in others it is more difficult or even impossible. The favourite and easiest
method is by means of CUTTINGS, that is, the planting of cut branches in water,
sand, or earth, in which they take root (Pelargonia, Tradescantias, Fuchsias,
Willows, Many plants may be propagated from even a single leaf or portion
etc.).
of a instance, is usually the case with Begonias.
leaf, as, for In other cases the
leaves, while still on the parent plant, have the power to produce adventitious
buds, and, in this way, give rise to new plants (see Vegetative Reproduction).
Even from roots or pieces of roots it is also possible to propagate some few plants.
An example of this is afforded by Ipecacuanha, whose roots are cut in pieces and
then sown like seeds. The Dandelion possesses the same capability of developing
from small portions of the root, and to this peculiarity is due the difficulty with
which it is destroyed.
In GRAFTING or BUDDING, cuttings from one plant are inserted in another, so that
they grow together to form physiologically one plant. The two parts stand in cor-
relation to one another, for a twig, which if planted in the soil would have developed
roots, when grafted on another plant forms no roots but enters into an intimate
relation with the stock. It adopts the roots of the latter and its buds may be said to
be adopted by the stock and no new organs are formed. When the affinity between
graft and stock is a distant one, it appears from LINDEMUTH'S observations that roots
may be formed from the graft even in the air. The union is accomplished by means
of a callus (p. 151), formed by both the scion and the adopted stock. Vessels and
sieve-tubes afterwards develop in the callus, and so join together the similarly
functioning elements of both parts. Such an organic union is only possible
between very nearly related plants, thus, for example, of the Amygdalaceae, the
Plum, Peach, Almond, Apricot, may readily be grafted one upon the other, or of
the Pomaceae, the Apple with the Quince but not the Apple with the Plum, nor
;

(as has been asserted) with the Oak.

252 BOTANY

In spite of the physiological union between the old stock and the newly
formed growth, from a morphological standpoint they lead an altogether separate
and distinct existence. They may, however, exert an influence on each other ;

thus annual plants grafted on biennial or perennial stocks attain an extended

period of existence. In its structural character, forms of tissues, mode of second-
ary growth, formation of bark, etc., each maintains its own individuality. In
special cases it has been affirmed that they do mutually exert, morphologically,
a modifying effect upon each other (graft-hybrids). This could only result from
fusion of the protoplasts of stock and graft in the callus (Fig. 247).
In practice several different methods of inserting grafts are in use, but only

Fie. 215. -Different modes of grafting ; /, Crown grafting ; //, splice grafting ; ///, Imrt grafting ;

IT, stock : K, scion.

the more important can be mentioned here. GRAFTING is the union of a shoot
with a young and approximately equally-developed wild stock. Both are cut
obliquely with a clean surface, placed together, and the junction protected from
the entrance of water and fungi by means of grafting wax (Fig. 215, II. ).
Cleft or tongue grafting is the insertion of weaker shoots in a stronger stock.
Several shoots are usually placed in the cut stem of the stock, care being taken
that the cambial region of the different portions are in contact, that the cortex of
the shoots is in contact with that of the stock. In other methods of grafting the
cut end of the shoot is split longitudinally and the cut shoot inserted in the

periphery, or a graft may be inserted in the cortex or in the side of the stock. In
grafting in the cortex the flatly-cut shoot is inserted in the space cut between the
bark and the splint wood (Fig. 215, /.). In lateral grafting, the shoot, after being
cut down, is wedged into a lateral incision in the stock.
A special kind of grafting is known as BUDDING (Fig. 215, ///.). In this process
 PHYSIOLOGY 253

"
a bud (" eye") and not a twig is inserted under the bark of the stock. The "eye
is attached to a shield-shaped piece of bark, which is easily separated from
left

the wood when the plants contain sap. The bark of the stock is opened by a
T-shaped cut, the "eye" inserted, and the whole tightly covered. Occasionally
some of the wood may be detached with the shield-shaped piece of bark (budding
with a woody shield). In the case of sprouting buds, the budding is made in
spring in dormant buds, which will sprout next year, in summer.
;

The Phase of Elongation

For the performance of their proper functions, the embryonic

rudiments of the organs must unfold and enlarge and assume their
characteristic appearance. This subsequent enlargement of the
embryonic organs of plants is accomplished in a peculiar and
economical manner. While the organs of animals increase in size
only by a corresponding increase of organic constructive material and
by the formation of new cells rich in protoplasm, and thus require
for theirgrowth large supplies of food substance, plants attain the
chief part of their enlargement by the absorption ofwater that is,
by the incorporation of an inorganic substance whichis most abund-

antly supplied to them from without, and to obtain which no

internal nutritive processes are first necessary.
The absorption of water by living cells does not take place with the same
rapidity and without interruption as in the case of porous bodies. Before the cells
can take up additional water they must enlarge by actual processes of growth.
The water, penetrating the young cells by imbibition or by the force of osmotic
pressure, is uniformly distributed through the protoplasm, which fills the cell ;

when the protoplasm already abundantly supplied with water, it is accumulated

is

in vacuoles (Fig. 58), As the vacuoles contain also organic and inorganic matter
in solution, they exert an attractive force and give rise to further absorption of
water. The sap of the vacuoles would, in turn, soon be diluted and its attractive
force diminished, were it not that the regulative activity of the protoplasm soon

provides for a corresponding increase of the dissolved salts, so that the concentra-
tion and attractive force of the sap are continually being restored or even increased.
The separate vacuoles thus enlarged ultimately flow together into one large sap-
cavity in the middle of the cell.

In this process the volume of the cell

may increase a hundred-
fold (in the Characeae two thousand-fold)
the internodal cells of
without any marked increase in the amount of protoplasm. The
enlargement of the cell has been almost wholly produced by the
increased volume of water in the sap cavity, which, to distinguish it
from the "nutrient water," "imbibition water," and "constitution
water" of the plant, may be designated "distension water."
As is observed in a whole series of vital phenomena, the rate
of distension of the walls with the distension water is not uniform,
but BEGINS SLOWLY, INCREASES TO A MAXIMUM RAPIDITY, AND
THEN, GRADUALLY DIMINISHING, ALTOGETHER CEASES. As all the
254 BOTANY PAUT i

cells of equal age in an organ go through this process of distension

at the same time, the phenomena of increase and decrease in the rate
of growth are apparent in the growth of the organ, and give rise to
GRAND PERIODS OF GROWTH. Minor periods, or fluctuations in the
rate of growth, occurring within the grand periods, are due to irregu-
larities in the swelling of the cells, occasioned by change of tempera-

ture, light, and other influences operative on growth, while the causes
5
of other abrupt changes in the rate of growth are still unknown ( ).
The large amount of water absorbed by the growing organ in the process of
elongation does not lessen its rigidity, but, on the contrary, it is to the turgor
thus maintained that the rigidity is due (cf. p. 178). Osmotic pressure seems
also to take an important part in the growth of the cell wall itself. Cells in whirh
the turgor is destroyed by a decrease in the water-supply exhibit no growth of
their cell walls;
it is thus evident that the distension of the cell walls is physically

essential for their surface-growth. This distension is in itself, however, by no

means the cause of their growth the internal physiological conditions of the
;

growth of the cell walls are dependent upon the activity of the living protoplasm.
Without the concurrent action of the protoplasm, there is no growth in even the
most distended cell wall on the other hand, active growth of the cell wall may
;

take place with the existence of only a small degree of turgor tension. A correspon-
dence between the turgor tension of the cell walls and the amount of growth cannot
under these conditions be expected, nor can, on the other hand, the conclusion be
drawn that turgor tension is inoperative in the processes of growth. The import-
ance of the turgor tensipn is variously estimated, according to whether the growth
of the cell wall is regarded as resulting from the interpolation of new particles of
constructive material between the already existing particles of the cell wall
substance (INTUSSUSCEPTION) or to the plastic (i.e. inelastic, not resuming its

original position) expansion of the distended cell wall. In the latter case the
growing membrane would continually become thinner, and require to be strengthened
by the deposition of new layers upon it (APPOSITION). Both processes, which may
occur together, probably take part in the growth of cell walls. The necessity of a
certain amount of turgor, if growth is to result from plastic stretching, is self-
evident ;
the stretching of the wall by the internal tension, though facilitating the
introduction of the new particles in growth by intussusception, is, however, not so
indispensable in this case.
The assumption of a growth by intussusception is intimately related to the views
held on the finer (or so-called molecular) structure of organic substances. The power
of swelling in water, which may even lead to complete solution, exhibited by organic
substances, shows that the water of imbibition does not merely penetrate into pre-
existing capillary spaces, but makes a passage for itself by separating the solid
particles from one another. It is further evident that these particles must be of
minute (molecular) size. The intimate penetration of the water is the expression of
a powerful molecular attraction, which is capable of exerting an enormous force it ;

is rendered
possible by the peculiar molecular construction of organic substances, the
cohesion of which is only gradually overcome by the water present in excess. The
arrangement of the particles has been pictured as resembling a network or a honey-
comb, while the frequent occurrence of double refraction as an optical property of
organic substances has been explained as due to the crystalline structure and definite
arrangement of the groups of molecules (micellae of NAEGELI) or to the relations of
tension in a colloidal honeycomb-like system (BiJTSCHLi). The new cellulose particles
 PHYSIOLOGY 255

would penetrate into the cell membrane, as particles of colouring matter may be
introduced with the imbibition water into a colloid organic substance C 86 ).

The process has so far been considered only in

of elongation
relation to the single cell, preparatory to the consideration of the

phenomena presented by the growth of multicellular organs. With

regard to this it is to be noted that the intensity of growth in two
organs of equal total growth is greater the shorter the growing zone
is. BUCHNER observed in fungal hyphae and pollen tubes in which
growth is restricted to just behind the apex that its amount per unit
of time was 220 per cent as compared with i-1 per cent in the
majority of the higher plants.
The operations of growth in plant organs proceed very slowly ;
so

Fit;. 21(5. .Simple and .self-reyisterinj; auxanometer. For description see text.

slowly as to be, in general, imperceptible. Only some fungal hyphae

and the stamens of many Gramineae grow so rapidly that their elonga-
tion is evident, even to the naked eye. The hyphae of Dictyophora
grow in length to the extent of 5 mm. per minute (A. MOLLER) and
according to ASKENASY, an increase in length of 1'8 mm. a minute
has been observed in the stamens of Triticum (Wheat). This approxi-
mately corresponds to the rate of movement of the minute-hand of a
watch. In comparison with these, the next known most rapidly
growing organ is the leaf-sheath of the Banana, which shows an elonga-
tion of I'l mm., and a Bamboo shoot, an increase in length of
0'6 mm. per minute; a strong shoot of Cucurbita grows O'09-O'l mm.
per minute, while most other plants, even under favourable circum-
stances, attain but a small rate of elongation (O'OOS mm. and less
67
per minute) ( ).
256 BOTANY PART I

In order to measure the growth in length of a plant, it is customary to magnify

in some way the actual elongation for more convenient observation. This may be
effected by means of a microscope, which magnifies the rate of growth corre-

spondingly with the distance grown. For large objects, the most convenient and
usual method of determining the rate of growth is by means of an AUXANOMKTKU.
The principle of all auxanometers, however they may differ in construction, is the
same, and is based upon the magnification of
the rate of growth by means of a lever with a
long and short arm. In Fig. 216, at the left, a
simple form of auxanometer is shown. The
thread fastened to the top of the plant to be
observed is passed over the movable pulley (r),
and held taut by the weight (</), which should
not be so heavy as to exert any strain on the
plant. To the pulley there is attached a slender
pointer (Z), which is twenty times as long as the
radius of the pulley, and this indicates on the
scale (S) the rapidity of the growth, magnified

twenty-fold. By a growth in the length of the

plant-stem of mm., the pointer would accord-
ingly register 4 mm.
Self-registering auxanometers are also used,
especially in making extended observations. In
Fig. 216, at the right, is shown one of simple
construction. The radius of the wheel () corre-

sponds to the long arm, and the radius of the

small wheel (r) to the short arm of the lever, in
the preceding apparatus. Any movement of the
wheel, induced by the elongation of the shoot,
and the consequent descent of the weight (G), is
recorded on the revolving drum ((7) by the pointer
attached to the weight (Z), which is, in turn,
balanced by the counterweight ( W}. The drum
Fio. 217. Unequal growth of different j s covered with smoked paper, and kept in rota-

regtons of the root-tip of Vicia Fa> M .

[m ,
the clock _ work {f7)> If the drum fe 8et
/, The root-tip divided by marking . . ,

with india-ink into 10 zones. e;u-h *> that it rotates on its axis every hour,
<>nct>

l mm. long. 17, The same root after the perpendicular distances between the tracings
twenty-two hours by the unequal
; on the drum will indicate the proportional hourly
growth of the different zones the grow ^i,
lines have become separated by mi- , .

equal distances. (After ACHS.) periods in the growth of an organ,

The grand
due to the internal causes, are clearly shown by
such self-registering auxanometers by the gradual increase and final decrease in
the perpendicular distances representing the increment of growth. STREHL found
the daily growth in length of a root of Lupine, expressed in tenths of millimetres,
to be o. For the
58, 70, 92, 97, 165, 192, 158, 137, 122, 83, 91, 59, 25, 25, 8, 2,
:

first internode of the stem, growing in the dark, the daily growth observed *aa
:

8, 9, II, 12, 35, 43, 41, 50, 51, 52, 65, 54, 43, 37, 28, 18, 6, 2, o.
The grand periods of growth, that is, the gradual increase from zero to a maximum,
and the succeeding decrease to zero again, are, however, not evident throughout the
a small portion of a root is
whole of a root ; during the growth in length only
actually, at one time, in process of elongation.
In roots of land plants the growing
SECT, ii PHYSIOLOGY 257

region extends over only about one centimetre of the extreme tip, often indeed over
only centimetre, while all the rest of the root has already completed its growth in
length. The length of the growing region is influenced by such external conditions
as mechanical hindrance, cold, warmth, dryness, etc., and exhibits consequent
variations. That the grand period is exhibited by this short growing region may
be made by marking oft' with india-ink, near the tip of a root, narrow zones
clear
of equal width, which would thus also be made up of cells of nearly equal size. In
Fig. 217 /, is shown a germinating Bean (Vicia Faba), whose root-tip has been
marked in this way Fig. 217 //, represents the same root after twenty-two hours
;

of growth. The marks have become separated by the elongation of the different
zones, but in different degrees, according to their position. The greatest elonga-
tion is shown by the transverse zone 3 ; from there the growth in length decreases
towards the younger zones (2 and 1), as well as towards the older (4 to 10). This
peculiar distribution of growth is but the result of the grand periods of growth of
the cells in zones of different ages ( 68 ). In the millimetre-broad zones of a root of
Vicia Faba SACHS found, after twenty-four hours, that the increase in growth,
expressed in tenth-millimetres, was as follows :

Zones: I., II., III., IV., V., VI., VII., VIII., IX., X., XI.
Increase :
15, 58, 82, 35, 16, 13, 5, 3, 2, i, o.
The elongating region much
longer than in roots, and
in shoot axes is generally
usually extends over several centimetres, in special cases even over 50 or more
centimetres. The distribution of the increase corresponds in stems, as in roots, with
the grand periods of, growth of the cells. Even by INTERCALARY GROWTH, where
the region of elongation is not confined to the apex but occurs in any part of the
organ, generally at its base (leaves and flower-stalks of many Monocotyledons),
grand periods of growth are also apparent. A shoot of Phascolus multiflor'ns which
was divided, from the tip downwards, into transverse zones 3*5 mm. broad, showed
in forty hours, according to SACHS,
in zones I., II., III., IV., V., VI., VII., VIII., IX., X.,
:
XL, XII.
an increase of 20, 25, 45, 65, 55, 30, 18, 10, 10, 5, 5, 5
tenth-millimetres.
This periodicity in the growth in length occurs even when the external influences
affecting growth remain constant, and is determined by internal causes alone.
Distinct periods of growth separated by an interval of time occur, according to
MIYAKE, in the scapes of the Dandelion, the first period in relation to the develop-
ment of the flowers, the second to that of the fruits. A similar behaviour is found
in other organs whose function after a time becomes altered (flower or fruit stalks
in Linana cymbalaria, and Arachis hypogaca (p. 285), and floral envelopes which
later protect the fruits in other cases).

External Influences upon Growth ( 69 ). External factors often

take an active part in the process of elongation, either as retarding
or accelerating influences. As growth is itself a vital action, it is
affected by any stimulus acting upon the protoplasm ; on the other
hand, as it is also a physical function, it is modified by purely physical
influences. Growth
is particularly dependent upon temperature,
light, moisture, the supply of oxygen and other substances, and the
existence of internal pressure and tension. It is also influenced by

injuries.
The INFLUENCE OF TEMPERATURE is manifested by the complete
s
258 BOTANY PAIST i

cessation of growth at a temperature less than or higher than

40-50. Between the MINIMUM and MAXIM CM temperatures, at which
growth ceases, there lies an OPTIMUM temperature (p. 176), at which
the rate of growth is greatest. This optimum temperature usually
lies between 22 and 37 C. The three CARDINAL POINTS OF
TEMPERATURE here given include a wide range, as they vary for
different species and even for individual plants of the same species,
and for their several vital processes. In tropical plants the minimum
temperature may be as high as + 10 C., while those of higher
latitudes, where the first plants of spring often grow through a
covering of snow, as well as those of the higher Alps and polar
regions, grow vigorously at a temperature but little above zero. In
like manner, the optimum and maximum temperatures show great
variation in different species of plants. Thus some Algae and Bacteria
grow in hot springs at a temperature of 80 C., which would be
at once fatal to other plants. The optimum does not usually lie in
the middle between the minimum and maximum, but is nearer the
maximum. (On the dependence of the cardinal points on external
conditions cf. p. 176.)
A certain amount of variation in the temperature favours the germination of
seeds and the unfolding of young shoots more than exposure to any constant
temperature. This is due to the optima for the different processes concerned lying
70
at different temperatures ( ).

The INFLUENCE OF LIGHT makes itself felt in a different manner

from changes of temperature. Light as a general rule retards
growth. This is apparent from observations on stems and roots
grown in the dark, and is also true in regard to the growth of leaves,
if the disturbing effects resulting from long-continued darkness be

disregarded. Too great an intensity of light causes a cessation of

the growth of an organ, while feeble illumination or darkness increases
it. The darkness upon the growth of plants is, however,
effect of

differently manifested according to its duration, whether it be

continuous, or interrupted, as in the changes of night and day.
Long-continued darkness produces an abnormal growth, in that the
growth of certain organs is unduly favoured, and of others greatly
retarded, so that a plant grown altogether in the dark presents an
abnormal appearance. The stems of Dicotyledons, in such case,
become unusually elongated, also soft and white in colour. The leaf-
blades are small and of a bright yellow colour, and remain for a long
time folded in the bud (Fig. 218, E). A plant grown under such con-
"
ditions is spoken of as etiolated."-

This diminution in the size of the leaf-blades and the elongation of the stem
(and leaf-stalks) are not manifested by all plants, nor under all circumstances. The
stems, for instance, of certain Cacti are much shorter when grown in the dark than
in the light, and their flattened shoots remain cylindrical. Similarly, the leaves of
 PHYSIOLOGY 259

varieties of the Beet (Beta) grow as large, or even larger, in tin- dark than in the

light ;
this is also true, under conditions favourable to nutrition, of the leaves of
other plants (Cucurbita). In the shade of a forest leaves often become larger than
in full daylight. are then proportionally thinner, and the palisade cells
They
which, in leaves fully exposed to the light, are in close contact, become pointed
below, and thus leave large intercellular spaces between
them. In this way the modifying influence of light of
diminished intensity is apparent in the internal structure
of such shade : leaves. Flowers, however, if sufficient
constructive material be provided by the assimilating
leaves, develop, according to SACHS' observations, as well
in the dark as in the sunlight, except that they are some-
times paler in colour. If, however, the assimilatory

activity of the green leaves be reduced or destroyed by

depriving them of light, many plants, as VOCHTING
found, form only inconspicuous or cleistogamous flowers ;

as (JOKBEL shows, the same result may follow when the

nutrition of the flowers is interfered with by poor soil or
the excessive development of the vegetative organs.
The tissues of etiolated stems and leaf-stalks are richer
in water and thinner- walled than in normally growing

plants. Even the roots of such plants are often found

to be less strongly developed. The supply of reserve
material at the disposal of plants growing in the dark
is utilised, with the assistance of an unusual amount

of distension water, in the elongation of the axis. This

elongation of the shoot axis, resulting from growth in
darkness, is of especial value in the development of
young plants from underground tubers, rhizomes, and
seeds. When the leaves must themselves reach the light

by their own elongation, as in many Monocotyledons,

they act just as do the stems of Dicotyledons, and attain
an abnormal length in the dark.

From what has already been said it would

seem that plants must grow more rapidly during
the night than day, and this is actually the case
where other conditions affecting growth remain Fl - ->18 -~ Two s edlin s s of
Sinapis alba, of equal age
the same by night as by day. A too low K, grown in the dark,
;

temperature during the night may, however, etiolated N, grown in or- ;

dinary daylight, normal.

completely counteract the accelerating influence The roots bear root-liaiis.
of darkness upon the growth.
.Just as the rays of light of different
wave-length and refrangibility
were found to be of different value in the process of assimilation, so
growth is by no means equal in differently coloured light. IT IS TO
THE STRONGLY REFRACTIVE SO-CALLED CHEMICAL RAYS THAT THE
INFLUENCE OF LIGHT ON GROWTH is DUE; the red-yellow end of the
spectrum acts upon many plants in the same manner as darkness.
Kadium and the Kontgen rays tend according to M. KORNICKE to
arrest the processes of growth and development ( 71 ).
260 BOTANY PART i

MOISTURE exerts a twofold influence upon growth. It acts as a

stimulus, and also, by diminishing transpiration, increases turgidity.

Plants in damp situations are usually larger than those grown in dry places,
and in fact may differ from them in their whole habit and mode of growth. Direct
contact with water seems frequently to exert a special influence upon the external
form of plants. Amphibious plants, that is, such as are capable of living both
upon land and in water, often assume in water an entirely different form from that
which they possess This variation of form is particularly manifested in the
in air.

leaves, which, so long as they grow in water, are frequently linear and sessile or
finely dissected, while in the air their leaf-blades are much broader and provided
with petioles (cf. Fig. 35). According to M'CALLUM, Proserpinaca palustris
forms dissected leaves even in a saturated atmosphere, while in salt solution it
develops leaves with a flat lamina as in the air. The leaf-stalks and internodes also
often exhibit a very different form in air and water, and undergo the same abnormal
elongation as in darkness. This is especially noticeable in submerged water plants,
whose organs must be brought to the surface of the water (young stems and leaf-
stalk of Trapa natans, stem of Hippuris, leaf-stalk of Nymphaea, Nuphar, Hy<lr<>-
charis). Such plants are enabled by this power of elongating their stems or lent
stalks to adapt themselves to the depth of the water, remaining short in shallow
water and becoming very long in deep water.
The great importance of free OXYGEN has already been alluded to in connection
with respiration (p. 240). Without gaseous or dissolved oxygen in its immediate
environment the growth of a plant entirely ceases, at least in the case of aerobionts.
MECHANICAL INFLUENCE. Pressure and traction exert a purely mechanical
influence upon growth, and also act at the same time as stimuli upon it. External

pressure at first retards growth it then, however, acccording to PFEFFEK, stimu-

;

lates the protoplasm and occasions the distension of the elastic cell walls, and

frequently also an increase of turgor. As a consequence of this increased turgor,

the counter-resistance to the external pressure is intensified. If the resistance of
the body exerting the pressure cannot be overcome, the plasticity of the cell walls
renders possible a most intimate contact with it thus, for instance, roots and root-
;

hairs which penetrate a narrow cavity so completely fill it that they seem to have
been poured into it in a fluid state. It would be natural to suppose that the effect
of such a tractive force as a pull would accelerate growth in length by aiding and
sustaining turgor expansion. But the regulative control exercised by the protoplasm
over the processes of growth is such that mechanical strain, as HEGLER has shown,
first upon growth to retard it (except in the maximum of the grand periods),
acts
but then causes an acceleration of even 20 per cent. According to BALL, a
constrained position may induce great thickening of the walls on particular sides
(e.g. the convex side) of an organ.
Rarefaction of the air, chemical stimuli, and internal states may exert consider-
able influence on growth. In this way the formation and development of the repro-
ductive organs, or the assumption of a twining habit may give rise to a striking
elongation of the internodes together with a reduction in the size of the leaves.
According to TOWNSEND, slight wounds accelerate growth, while more serious
72
ones retard it ( ).

The Internal Development of the Organs

The
internal development of the organs is only completed after
they have finished their elongation and attained their ultimate size.
 PHYSIOLOGY 261

They are then first enabled to fully exercise their special function.
To this end the cavities of certain cells usually become more or less
fused, and the cell walls thickened, often in a peculiar and character-
istic manner (p. 63).

Periodicity in Development and Duration of Life

The periodically recurring changes in the determinative external

influences, especially in light and temperature, occasioned by the
alternations of day and night and of the seasons, cause corresponding
periodical variations in thegrowth of plants. These variations do
not follow passively every change in the condition of the external
influences. On the contrary, the internal vital processes of plants so
accommodate themselves to a regular periodicity that they continue
for a time their customary mode of growth, independently of any
external change. The nightly increase of growth, which is especially
noticeable after midnight in the curve of growth, and the retardation
of growth, specially marked after mid-day, will continue to be
exhibited for some time in prolonged darkness when the temperature
remains constant. Thus under these conditions Helianthus tuberosus has
been observed to continue its regular DAILY PERIODS for two weeks,
affording an example of an after effect of the usual rhythm which
suggests a comparison with the faculty of memory, and will be further
considered below. According to SEMON, the usual periodicity may
even be transmitted to the descendants of a plant.
Still greater is the influence exerted on the life of plants by the
alternation of winter and summer, which in the plants of the colder
zones has rendered necessary a well-marked winter rest. This is not
in reality an absolute rest ; for although the outwardly visible pro-
cesses of development and growth stand still, the internal vital
processes, although retarded, never altogether cease.
The ANNUAL PERIODS of growth occasioned by climatic changes, which are
rendered so noticeable by the falling of the leaves in the autumn, and the develop-
ment of new shoots and leaves in the spring, have stamped themselves so indelibly
upon the life of the trees and shrubs of the temperate zones, that, when cultivated
in tropical lands where other plants are green and blossom and bear fruit through-
out the year, they continue to lose their leaves and pass for a short time at least
into a stage of rest. The Oak, Beech, Apple, and Pear retain their resting period
in the sub -tropical climate of Madeira, while, under uniformly favourable
conditions in the mountain regions of Java, the periodicity may be disturbed in
particular individuals. This even occurs in the several brandies of the same tree
which may then bear leafy and leafless boughs at the same time (Oaks. Magnolias,
Fruit and Almond trees, together with some endemic species). Other trees again
gradually accustom themselves to the new conditions, as the Peach, for instance,
which in Ceylon has become an evergreen tree. The Peach is reported to produce
flowers and fruit throughout the entire year while the Cherry, like many other
;

trees of the temperate zone, ceases altogether to bear flowers in tropical climates.
262 BOTANY PAI:T i

Since in these tropical localities endemic species may exhibit well-marked periods
of rest and of renewed vegetative activity, it is clear that such periodicity is not
merely induced and regulated by external influences. Its cause must rather be
sought in an autonomous, rhythmic course of the vital process itself. Although to
so many plants winter is the season of rest and cessation from growth, other

plants, e.g. certain Lichens and Mosses, seem to find in the warmer days of winter
the most favourable conditions of vegetation and in summer, on the contrary,
;

either do not grow at all or

only very little. Similarly, many spring plants attain
their highest development, not in summer, but during the variable weather of
March and April, and, for the most part, they have entered upon their period of
rest when the summer vegetation is just awakening.
In countries where there are alternate rainy and dry periods, the latter
generally corresponds to the period of vegetative rest.
Favourable conditions may shorten the resting period, especially if tin-

organisms havepreviously been exposed to the influence of frost, drought

(withering), or, as JOHANNSEN discovered, to ether vapour, etc. This premature
awakening does not, however, result equally readily at all periods of the resting time.
It occurs best in the later portion of the latter, just before the normal time of

awakening, and almost equally well in the earlier portion just after arrest has taken
place. Potato tubers and bulbs in some cases do not completely enter into a
resting state before their development is awakened. In the intervening period of
complete rest such attempts are usually without result. These facts must l>e
considered in the forcing of plants, as must also the fact of the temperature
optima differing for the several developmental processes, for some of which tin \
are relatively low (
7:i
).

DURATION OF PLANT LIFE. The life of a plant, during the whole

of its development, from its germination to
death, is dependent upon its

external and internal conditions. In the case of the lower vegetable

organisms, such as Algae, Fungi, and Bacteria, their whole existence
may be completed within a few days or even hours, and indeed some
of the higher herbaceous plants last only for a few weeks, while the
persistent shrubs and trees, on the other hand, may live for thou-
sands of years.

After the formation of the seeds, there occurs in many plants a cessation of
their developmental processes, and such a complete exhaustion of vitality that
death ensues. Such an organic termination of the period of life occurs in our
annual summer plants, which, according to KLKBS, can be made perennial by
artificiallypreventing their reproduction. It also takes place with plants in
which the preparatory processes for the formation of fruit have extended for two
or more years, as in biennial plants in the case of the 10- to 40- year-old Agave,
which, after the formation of its stately inflorescence, dies of exhaustion, and in
some Palms (Corypha umbraculifera). In plants, on the other hand, which in
addition to the production of flowers and fruit accumulate also a reserve of organic
substance, and, with their reproductive organs, form also new growing points, life
does not cease with the production of the seeds. Such plants possess within
themselves the power of unlimited life, the duration of which may only be
terminated by unfavourable external conditions, the ravages of parasites, injuries
from wind, and other causes.
The longevity of trees having an historical interest is naturally best known
 PHYSIOLOGY 263

and most celebrated, although, no doubt, the age of many other trees, still living,
dates back far beyond historical times. The celebrated Lime of Neustadt in Wurtem-
berg is nearly 700 years old. Another Lime 257 in. in circumference had 815
annual rings, and the age of a Yew in Braburn (Kent) which is 18 m. in circumference
is estimated at 2880 years. A stem of a Sequoia in the Berlin Museum has, with
1360 annual rings, a diameter of 47 m., from which an idea can te formed of the
age of those trees which have attained a diameter of 16 m. An Adansonia at
Cape Verde, whose stem is 8-9 m. in diameter, and a Water Cypress ( Taxodium
mexieantiM) near Oaxaca, Mexico, are also well-known examples of old trees. Of
an equally astonishing age must have been the celebrated Dragon tree of Orotava,
which was overturned in a storm in 1868, and afterwards destroyed by fire. The
lower plants also may attain a great age the apically growing mosses of the
;

calcined Gymnostomum clumps, and the stems of the Sphagnaceae, metre-deep in

a peat-bog, must certainly continue to live for many centuries ( 74 ).

In thus referring to the 'ages of these giant plants, it must not

be understood that all the cells remain living for so long a time, but
rather that new organs and tissues are developed, which continue
the life of the whole organism. All that is actually visible of a
thousand-year-old Oak is at most but a few years old. The older
parts are dead, and are either concealed within the tree, as the pith
and wood, or have been discarded like the primary cortex. The cells
of the original growing point have alone remained the whole time
alive. They continue their growth and cell division so long as the
tree exists ; while the somatic cells of the permanent tissue arising
from them, and destined for particular purposes, all lose their vitality
after a longer or shorter performance of their functions.

The cells of the root-hairs often live for only a few days ;
the same is also true
of the glandular cells and trichomes of stems and leaves. The wood and bast fibres,
as also the sclerenchymatous cells, lose their living protoplasiu after a short time.
Entire organs of long-lived plants have frequently but a short existence the sepals, ;

petals, and stamens, for example. The foliage leaves, also, of deciduous trees live
only a few summer months and then are discarded. The leaves even of evergreen
'

plants continue living but a few years, before they too fall off. Small twigs,
especially of Conifers, are also subject to the same fate. The cells of the medullary
rays afford the best examples of long-lived cells constituting permanent tissues.
In many trees, as in the Beech, living medullary ray cells over a hundred years
old have been found, although, for the most part, they live only about fifty years.

V. The Phenomena of Movement

In every living organism there is constantly occurring in the

course of the metabolic processes an active movement and transposi-
tion of substance. As these movements are for the most part
molecular they are generally imperceptible ; but that they actually
take place is demonstrated with absolute certainty by the local
accumulation and diminution of substance.
There are also other forms of movement which play an important
264 BOTANY PART I

part in the physiology of every organism, and on which its vital

processes are to, a large extent dependent these are the movements
:

due to heat and the related conditions of vibration resulting from

light, electricity, etc.
Apart from the movements of this class, which may take place
within organisms which, externally, are apparently at rest, there occur
also in plants actual CHANGES IN POSITION, externally noticeable but

usually of gradual operation ; in special cases they may involve

rapid motion. These movements may be carried on either by the
whole plant or by single organs.

Reference is here SPONTANEOUS MOVEMENT resulting from the

made only to the
and this should not be confused with the PASSIVE
activity of a plant organism itself,
movements due to externally operating mechanical agencies, such as water and
wind, which, although they have a certain importance for plant life, will not
be here considered.

PROTOPLASM itself is capable of different movements. Naked

protoplasmic bodies almost always show slow movements resulting in
a gradual change of position ; but cells enclosed by cell walls possess
also the power of INDEPENDENT LOCOMOTION, often indeed to a con-
siderable extent. Multicellular plants, however, as a rule ultimately
attach themselves, by means of roots or other organs, to the place of
germination, and so lose for ever their power of locomotion, except in
so far as it from growth. A gradual change in position due
results
to growth apparent in plants provided with rhizomes, the apical
is

extremities of which are continually growing forward, while the older

portions gradually die off. A yearly elongation of 5 cm. in the
apical growth of the rhizomes would result, in twenty years, in
moving the plant a distance of one metre from its original position.
A seedling of Ouscuta in its search for a host plant illustrates the
power of maintaining, for a time, a creeping movement over the
surface of the soil; a growing Caulerpa (Fig. 295) likewise exhibits in
the course of years a similar advancing movement. In addition to
these movements, occasioned by a growth in length, plants firmly
established in the soil possess also the power of changing the position
and direction of their organs by means of CURVATURE and ROTATION.
In this way the organs are brought into positions necessary or
advantageous for the performance of their function. By this means,
for example, the stems are directed upwards, the roots downwards ;

the upper sides of the leaves turned towards the light, climbing
plants and tendrils twined about a support, and the stems of
seedlings so bent that they break through the soil without injury to
the young leaves.
 PHYSIOLOGY 265

Movements of Naked Protoplasts and Single Cells

The creeping movements of naked protoplasts, such as are shown

by an amoeba or plasmodium, in the protrusion, from one or more
sides, of protuberances which ultimately draw after them the whole
protoplasmic body, or are themselves again drawn in (Fig. 219), are
distinguished as AMOEBOID MOVEMENTS. These movements resemble,
externally, the motion of a drop of some viscid fluid on a surface to
which it does not adhere, and are chiefly due to surface tension,
which the protoplasm can at different points increase or diminish, by
means of its quality of irritability. By means of irregular changes
of surface-tension similar amoeboid move-
ments are also exhibited by drops of life-
less such as drops of oil in soap
fluids,
solution ;drops of mercury in 20
or
per cent solution of potassium nitrate in
contact with crystals of potassium bi-
chromate.
In the SWIMMING MOVEMENTS BY
MEANS OF CILIA, on the contrary, the
whole protoplasmic body is not involved,
but it possesses special organs of motion
in the form of whip-like FLAGELLA or
CILIA. These may be one, two, four, or
FIG. 211>. AiiKtboid movement. The
more in number, and arranged in various arrows indicate the direction and
ways (Figs. 97, 98). They move very energy of the movement the
;

At the
crosses, the points at rest.
rapidly in the water and impart con- time being the principal movement
siderable velocity to the protoplast, often is from H to V, but at any moment

it at the same time a rotary move- it may be towards R or L, and so

giving
changei the direction of the course
ment. The minute swarm -spores of taken by the amoeba.
Fuligo varians traverse 1 mm. (sixty
times their own length) ina second, those of Ulw, O'lo mm., while
others move more slowly. The Vibrio of Cholera, one of the most
rapidly moving bacilli, takes 22 seconds to traverse a millimetre.
The ciliary movement is so regulated as to propel the protoplast in
a definite direction.

Gravity and light, certain substances in solution, and mechanical hindrances

are the principal influences which regulate the movements of free-swimming proto-

plasmic bodies and cells. The direction of the movements of the swarm-spores of
Algae are chiefly determined by the light. So long as they remain in darkness
they move through the water in all directions but as soon as they are illuminated
;

from one side only, a definite direction in their movements is perceptible. They
move either straight towards the light or turn directly away from its source.
Their retrogressive movements from the light occur either in case of too intense
illumination, or at a certain age, or through some unknown disturbing irritation.
The advantage of such HELIOTACTIC MOVEMENTS (phototactic) is at once apparent
266 I50TANY PAIIT i

when the part taken by the swarm-spore in the life of an Alga is considered. In
order to provide for the future nutrition of the stationary Alga into which it
afterwards develops, it must seek the light. If a point with suitable (that is, not
too intense and not too weak) illumination be attained, then the swarm-spore must
attach itself by the end which carries the cilia to do this it must turn itself from
:

the light towards a dark object. On the other hand, as the swarm-spores do not
come to rest at all in absolute darkness, but swim continuously until thoroughly
exhausted, the possibility of their attaching themselves in a spot devoid of light
where the new plant could not assimilate is excluded.
The swarm-spores of aquatic Fungi and spermatozoids, according to PFKFFEK'S
investigations, are chiefly influenced in their movements by the unequal distribu-
tion of dissolved substances in their environment (topotaxis). The crowding
together of Bacteria appears, from ROTHEUT'S experiments, not to be due to move-
ment towards an attracting stimulus but to their inability to leave a point of
optimal concentration reached by accident ; the decline in this concentration
arrests their movement and prevents their dispersal (phobotaxis). Minute traces
of free oxygen can be recognised by the influence exerted on the movement of
certain Bacteria ;
ENGELMAXN'S
bacterial method of demonstrating assimilation

depends on this According to their momentary requirements and their

fact.
sensitiveness to stimuli, such small organisms move either towards or away from
the points of highest concentration they are sensitive to the quantity as well
;

as the quality of the dissolved substances, and also to the osmotic effects of the
latter (osmotaxis) 73
( ).

As the CHEMOTACTIC MOVEMENTS spermatozoids approach the

result of similar
female sexual organs. PFEFFER has demonstrated that the spermatozoids of
Ferns are enticed into the long necks of the archegonia by means of malic acid :

while the archegonia of the Mosses attract the spermatozoids by a solution of cane-
sugar, and those of the Marchantiaceae by proteid substances. In such cases an
extremely small quantity of dissolved substance is often a sufficient stimulus to
call forth active chemotactic movements a O'OOl per cent solution of malic acid
;

suffices for the attraction of Fern spermatozoids. The movements of amoebae and
plasmodia are similarly induced by external influences. These naked protoplasts
live not only in water but also in moist substrata (plasmodia, amoebae), and seem
to possess the power of seeking out situations with more moisture, or of avoiding
them (before the formation of spores) their movements are also influenced by the
:

direction of currents in the water (rheotaxis). The term thigmotaxis or stereo-

taxis has been applied to the exceptional property of certain swarming cells, which
are stimulated by mechanical contact to remain attached to the object around which
they are swarming. This has been observed in spermatozoids of Fucaceae and in
Chromatium wcissii, a sulphur Bacterium. The fact that algal swarm-spores on a
hard rough substratum form stronger attaching organs than on a smooth surface is
related' in some degree to this. In cases where cells enclosed by cell walls (e.g.
Haematococcus phnnalis) swim freely about by means 'of cilia, the cilia spring
from the protoplasm and pierce the cell walls.
Diatoms and Desmids exhibit quite a different class of movements. The
Diatoms glide along, usually in a line with their longitudinal axes, and change
the direction of their movements by oscillatory motions. From the manner in
which small particles in their neighbourhood are set in motion, it was concluded
that special organs of motion probably protrude, like pseudopodia, through
openings in their hard silicified shell. Recently these motile organs have been
seen in some forms. According to O. MULLEU, the movement is set up by a
 PHYSIOLOGY 267

current of protoplasm, which bursts through the raphe this may become invested
;

with a mucilaginous sheath. Corresponding to the differences in the construction

of the membrane, differences in the motile apparatus are found in the group. The
cells of Desmidiaceae the substratum by mucilaginous
attach themselves to
excretions, and effect movements by local fluctuations in the
their peculiar
mucilaginous layer. The advancing movements of the filamentous Oscillarieae
and Spirulinae take place, according to CORUENS, in a mucilaginous sheath, but
their mechanism is as little understood as that of the slow movements of

Spirogyra.

The Movements of Protoplasm within Walled Cells

Although plants which are firmly attached and stationary exhibit

no such locomotory movements, the protoplasm within their cells
does possess a power of movement. Such internal protoplasmic
movements are especially active in the non-cellular Siphoneae, and in
non-septate Fungi in the elongated internodal cells of the Characeae,
in the hairs of many plants, as well as in the leaf-cells of some

aquatic plants.
The three following different forms of protoplasmic movement
within cell cavities may be distinguished CIRCULATION, ROTATION,
:

and ORIENTATION.
In the case of CIRCULATORY MOVEMENT the different currents of
protoplasm, although often quite close together, flow in different
directions in slender protoplasmic strands, which stretch from the
cell wall to the nucleus (cf. p. 58 and
Fig. 60).
In the ROTATORY MOVEMENT the protoplasm moves along the cell
wall in one direction only, dragging with it the nucleus and often
also the chlorophyll grains (cf. p. 58).

The cause of these movements, which may take different directions in adjoining
cells, and may also continue after the protoplasm has been drawn away from the
cell walls by plasmolysis (p. 180), is not yet understood. It is, however, known
that the continuance and activity of such protoplasmic movements, the existence
of which was first observed by CORTI in 1772, and later rediscovered by
TIIKVIRANUS in 1807, are dependent on factors which, in general, support and
promote the vital activity while the presence of free oxygen and proper
;

conditions of temperature seem to be particularly favourable to them. That the

movements in Nitella continued, according to KUHNE'S observations, for days and
weeks in the absence of oxygen is explained, according to HITTER, by the power
the Characeae possess of becoming facultative anaerobes. Those Bacteria which
are obligate anaerobes lose their power of motion in the presence of oxygen.
From the study of sections in the cells of which currents had been induced in
the protoplasm, by the injuries sustained in their preparation and by other
abnormal conditions, grave errors have arisen concerning the existence of such
protoplasmic movements in cells, in which under normal conditions they cannot
be observed. The presence of protoplasmic currents in a cell may, in fact,
indicate either an energetic vital activity, or, on the other hand, be merely a
symptom of a pathological or, at least, of an, excited condition of the proto-
7B
plasm ( ).
268 BOTANY PAKT i

The movements of orientation of the protoplasmic body are usually

so gradual as to be only recognisable through their operations. They
are induced by changes in the external influences, especially as
regards the intensity of the light, and result in producing a definite
position of the protoplasmic bodies, as, for example, the orientation
of the chlorophyll grains with regard to the light. Movements of

FIG. 221.
Streaming protoplasm
inthe hyphse of 7tfii :'v<> <;//
<'-

cans. ( After J. C. ARTHUR.)

this kind have been most frequently

observed in Algae, in submerged Duck-
weed (Lemna trisulca), in the prothallia
ofFerns and Mosses but similar move-
;

ments can also be observed in the higher

plants.
Kin. ~2'20. Varying positions taken by
the Chlorophyll grains in the cells of In the cells of the filamentous Alga Meso-
I*mna triscvlca in illumination of dif- carpus, the chloroplasts, in the form of a single
ferent intensity. T, in diffuse day-
* kte sus ded length-wise in each cell, turn
light; S, in direct sunlight; N, at \
u l (On thelr longitudinal axes according to the
night. The arrows indicate the direc-
tion of the light. (After STAHL.) direction and intensity of the light. In light of
moderate intensity, according to STAHL'S ob-
servations, they place themselves transversely to the source of light, so that
they are fully illuminated (transverse position) ; when, on the other hand, they
are exposed to direct sunlight, the chlorophyll plates are so turned that their
edges are directed towards the source of light (profile position). similar pro- A
tection of the chloroplasts against too intense light, and their direct exposure,
on the other hand, to moderate illumination, is accomplished, where they are of a
different form and more numerous, by their different disposition relatively to the cell
walls. In moderate light the chlorophyll bodies are crowded along the walls, which
 PHYSIOLOGY 269

are at right angles to the direction of the rays of light (Fig. 220 T). They quickly
pass over to the walls parallel to the rays of light, however, as soon as the light
becomes too intense, and so retreat as far as possible from its action (Fig. 220 S).
In darkness or in weak light the chloroplasts group themselves in still a third way
(Fig. 220 A"), the advantage of which is not altogether clear. Similar changes of
position may result from certain substances in solution in the surrounding water.
The form of the chlorophyll bodies themselves undergoes modification during
changes in their illumination ; in moderate light they become flattened, while in
light of greater intensity they are smaller and thicker.
As a special mode of protection against too intense light, the chloroplasts of the
Siphoneae and Diatomeae (and the same thing is observed in many plants) become
balled together in separate clumps. In correspondence with the changes in the
position of the chloroplasts, the colouring of green organs naturally becomes
modified. In direct sunshine they appear lighter, in diffused light a darker green.
The attention of SACHS was first called to the phenomena of the movements of the
chloroplasts by the accidental observation that the shadow of a thermometer was
represented in dark green on a leaf otherwise directly illuminated by the sun.
Wounds and cell-wall thickenings localised to one side of the cell likewise give
rise to orientation movements, as they occasion a crowding together on one side
of the nucleus and protoplasm.

Apeculiar mode of protoplasmic movement, which may be termed

STREAMING, occurs, according to ARTHUR, in the non-septate mycelium
of some Fungi. In it the whole mass of protoplasm, with its included
vacuoles, streams towards the end of the hypha, only the limiting
layer remaining at rest. After a longer or shorter interval a similar
streaming movement sets in in the opposite direction (cf. Fig. 221).
TERNETZ observed the same type of movement in the mycelium of Ascophauus
carneus, where the protoplasmic stream, with its vacuoles, makes its way through
the pores of the perforated transverse septa. In this latter case it was established
that the direction of streaming was determined by local differences in the supply
of water C77 ).

Movements producing Curvature

The movements of the organs of stationary plants, unicellular as

well as multicellular, are accomplished by means of curvatures. In an
organ that has grown in a straight line the longitudinal sides are all
of equal length ; in an organ that is curved, however, the concave
side is necessarily shorter than the convex side. When, accordingly,
the opposite sides of a pliable organ become of unequal length, the
organ must curve toward the shorter side (Fig. 175). Inequality in
the length of the opposite sides may result from various causes. A
curvature occurs if the length of one side remains constant, while the
opposite side becomes shorter or longer, and also from the unequal
elongation or contraction of both sides, or from the elongation of one
side and the contraction of the other.
Such curvatures most frequently occur in plants as a consequence
of UNEQUAL GROWTH. More rarely they are due to the different
 270 HOTANY I..VKT i

length of the opposite sides, resulting from unequal TURGOR TENSION.

A third source of curvature is found in the unequal amount of water
taken up by IMBIBITION, and the consequent unequal swelling of the
cell walls on the opposite sides of an organ. A
fourth cause is the
loss of water from certain cell-complexes, the latter being made to
contract in a definite direction owing to the cohesive force of the
water which remains in them.
Whenthe direction of the curvature is determined by the position
of the source of the stimulus causing it, the movement is termed tropic

(tropism) ; when no such relation is evident and the movement results

from the internal disposition of the structure, it is spoken of as nastic.

1. Movements dependent on Imbibition and Cohesion

As the cell Avails of actively living cells are always completely

saturated with imbibition water, hygroscopic curvatures are exhibited
only by dry or drying and, for the most part, dead tissues. The
hygroscopic movements in any case, however, are due to the physical
properties of the cell walls, and have no direct connection with the
vital processes, except in so far as the capacity of cell walls to swell
and take up large quantities of imbibition water is due to the proto-
plasm by which they were formed. The activity of the protoplasm
in the formation of the cell walls is likewise manifested in their
anatomical structure, in their stratification and striation, and in the
position of the pits.
The absorption of imbibition water by cell walls is accompanied
by an increase in their volume, and conversely the volume of the cell
walls is diminished by the evaporation of the imbibition water. Ac-
cordingly, whenever unequal amounts of water are held by the cell
walls on the different sides of an organ, either through unequal
absorption or evaporation, hygroscopic movements are produced,
which result in the curvature of the organs. In many cases the
organs of plants are especially adapted to such movements, by means
of which, in fact, important operations are often accomplished, as, for
example, the dehiscence of seed-vessels and the dissemination and
burial of seeds. Similar movements also occur, without special
purpose, however, as for example in dead branches of Conifers.

The rupture of ripe seed-vessels, as well as their dehiscence by the opening of

special apertures (Papaver, Lychnis, Antirrhinum, etc.), is a consequence of the
unequal contraction of the cell walls due to desiccation. At the same time, through
the sudden relaxation of the tension, the seeds are often shot out to a great dist.mn'
(Tricoccae, Geranium, etc.). In certain fruits not only curvatures but torsions ;m-
produced as the result of changes in the amount of water they contain, e.g. Erodiutn
gruinum (Fig. 222), Stipa pennata, A vena sterilis ; by means of these, in conjunc-
tion with their stiff barb-like hairs, the seeds bury themselves in the earth.
 PHYSIOLOGY 271

The variation in dampness of the air causes the pappus hairs of the
Cynareae (Compositae) to expand in dry and fold together in damp weather.
The opening or closing of the moss sporogonium is, in like manner, due to the
hygroscopic movements of the teeth of the peristome surrounding the mouth
of the capsule. In the case of the Equisetaceae the outer walls of the spores
themselves (the perinium) take the form of four arms, which, like elaters, are cap-
able of active movements, by means of which numbers of spores become massed

together before germinating, and the isola-

tion of the dioecious prothallia prevented

(Fig. 420E).
In order to call forth imbibition move-
ments the actual presence of liquid water
is not necessary for the cell walls have
;

the power of absorbing moisture from the

air. They are hygroscopic, and for this
reason the ensuing movements are also
often termed hygroscopic movements.

STEINBRINCK and KAMERLING

have distinguished the mechanisms
which depend on the cohesive power
of water from those depending on
imbibition. The cohesion mechan-
isms were previously confounded
with the latter, from which they
differ in that, even during the move-

ment, the cell walls remain saturated

with water. It is the lumen of the
cell which diminishes in size when
the loss of water, on which the
movement depends, occurs. The
cohesive of the diminishing
force
amount of water tends to pull the
thinner walls of the cell inwards
after it, and thus to approximate 2 Pai frnitrf J5rKw jrf MW.
A, in the dry condition, coiled; B, moist
other walls, which are strongly ., , ,., 1(11 iltl .;,.

thickened and exhibit a definite

arrangement. In this way an energetic shortening of the specialised
tissue is brought about which leads to alteration of form or to the
dehiscence of spaces enclosed by the tissue. The walls of anthers
and of the sporangia of the higher cryptogams afford examples of
such a method of dehiscence. The movements of the elaters of
Liverworts and Myxomycetes, which serve to distribute the spores,
as well as of the pappus of most, and the involucre of some, Com-
78
positae, are effected in the same way ( ).
Mechanisms dependent on imbibition and on cohesion may co-
operate in the movements of some plants.
272 BOTANY

2. Growth Curvatures

Movements from which curvatures result are, for the most part,
produced by the unequal growth of living organs. The unequal
growth is due, partly to internal causes which are still undetermined,
and partly to the operation of external influences. The movements
resulting in the first case are spontaneous, and are called AUTONOMIC
MOVEMENTS or NUTATIONS ; in the second case the movements are
the result of external stimuli, and are distinguished as IRRITABLE or
PARATONIC (AITIONOMIC) MOVEMENTS.
Autonomie Growth Curvatures (Nutations) are most plainly
apparent in young actively-growing organs, although nutations have
been shown to be exhibited by all growing plants, as their tips do
not grow forward in a straight line, but, instead, describe irregular
elliptical curves. These movements, which Darwin termed CIRCUM-
NUTATIONS, while often not perceptible to the eye, are very noticeable
is some special organs.

The unfolding of most leaf and flower buds, for example, is a nutation move-
ment which, in this instance, is induced by the more vigorous growth of the inner
side of the young leaves. The same unequal growth manifests itself most noticeably
in the leaves of Ferns and many Cycadeae. In the same manner, movements of
nutation are caused in lateral axes when growth is more energetic on either the
upper side (EPINASTY) or on the lower side (HYPONASTY). Epinastic curvatures are
often greatly increased when poisonous gases are present in the air, and may also be
" load-curvature "
started by the organ's own weight. This is termed by WIKSNER.
The stems of many seedlings are, on their emergence from the seeds, strongly
curved. By the nutation of the shoots of Ampclopsis quinquefolia a curvature
isproduced which continuously advances with the increased growth .so that, by
;

means of its hooked extremity, a shoot is better enabled to seek out and cling to
a support. When the unequal growth is not confined to one side, but occurs
alternately on different sides of an organ, the nutations which result seem even
more remarkable. Such movements are particularly apparent in the flower-stalk
of an Onion or of Yucca filamentosa, which, although finally erect, in a half-grown
state often curves over so that its tip touches the ground. This extreme curvature
is not, however, of long duration, and the flower-stalk soon becomes erect again
and bends in another direction. Thin and greatly elongated organs (e.g. tendrils)
must, from purely physical reasons, quickly respond to the effects of unequal
growth, If the line of greatest growth advances in a definite direction around the

stem, its apex will exhibit similar rotatory movements (REVOLVING NUTATION).
This form of nutation is characteristic of the tendrils and shoots of climbing plants,
and facilitates their coming in contact with a support. The so-called REVOLVING
NUTATION OF TWINING PLANTS is not, however, an AUTONOMIC MOVEMENT, and

will be considered later with the paratonic movements.

Paratonic (Aitionomie) Growth Curvatures. The paratonic move-

ments are of the very greatest importance to plant life, for through
their operations the organs of plants first assume such positions in
 PHYSIOLOGY '27:;

air,or water, or in the earth as are necessary for the performance of

their vital functions. A
green plant which spread its roots over the
surface and unfolded its leaves below ground could not exist, even
though all its members possessed the best anatomical structure.
Seeds are not always deposited in the soil with the embryonal stem
directed upwards and the radicle downwards, so that their different
organs can, merely by direct growth, attain at once their proper
position. A
gardener does not take the trouble to ascertain, in
sowing seed, if the end which produces the root is directed downwards
or the stem end upwards he knows that in
; any position the roots
grow into the ground and the stems push themselves above the
surface. Plants must accordingly have in themselves the power of
placing their organs in the positions best adapted to the conditions
of their environment. That is only possible, however, when the
externally operative forces and substances can so influence the
growth of a plant that it is constrained to take certain 'definite
directions.
The same external influences excite different organs to assume
quite different positions. Through the influence of gravity, the tap-
root grows directly downwards in the soil, while the lateral roots
take a more or less oblique direction. The main stem grows
perpendicularly upwards it, like the primary root, is ORTHOTROPIC.
;

The on the other hand, just as the secondary roots,

lateral branches,
assume an inclined position and are PLAGIOTROPIC. The apical ex-
tremities of shoots are constrained to seek the source of light ; the
leaves, on the contrary, under the same influence place their surfaces
transversely to the direction of the rays of light. The property
whereby an organ, when acted upon by external influences, assumes
different positions has been termed ANISOTROPY by SACHS. In
addition to the purely morphological structure of the plant body,
anisotropy also determines essentially its external form and appear-
79
ance, or what is termed the habit of the plant ( ).
That all these paratonic movements cannot result merely from
the physical action of external forces will be at once recognised if it be
taken into consideration that anisotropic but in other respects similar
organs are affected differently by the same influences, and that even
the same organs react differently at different ages ; moreover, the
external forces produce effects which bear no relation to their usual
physical and chemical operations. It will, on the contrary, be at once

apparent that the movements are rather the result of definite pro-
cesses of growth, arising from an irritability to stimuli induced by
external influences (cf. pp. 4, 174).
In order that external influences may produce such efl'ects, plants must be
sensitive to stimuli, that is, the stimuli must produce in them certain modifications
with which, in turn, certain definite vital actions are connected. The precise
manner in which an external influence produees an internal reaction within an
T
274 BOTANY PAHT i

organism is not at present known. In order that an external physical force can
operate as a stimulus, there must exist within the living substance definite
structures or organs which are influenced by it. Thus, for example, H.\r,r.i:i AMU
and NEMEC regard the pressure of starch grains (the position of which in the cell is
determined by their weight) on the more or less receptive parts of the limiting
layer of the protoplast as the arrangement for perceiving the direction of gravity
(statolith theory). The movements of growth occasioned by external stimuli are,
for the most part, movements in response to directive stimuli which lead to a
definite position of the organ, relatively to the direction of the operative influence.
The principal external stimuli that come into consideration are light, heat, gravity,
chemical influences (oxygen, nutritive substances, water, etc.), impact and
friction.
As the
points of greatest irritability in plants or their organs are often more or
lessremoved from the points where the effect of the stimulation is manifested, a
propagation of the stimulation must take place. Thus directive stimuli are
received by the apices of organs which do not themselves carry out the movement,
but transmit it to the regions in which the movement takes place. In root* tin
geotropic stimulus mainly perceived by the root tip ; in grass-seedlings the
is

stimulus of light perceived by the tip of the cotyledonary sheath ; in the

is

tentacles of Drosera the contact stimulus affects the glandular swelling at the tip
and induces the movement in the portions nearer the base. According to MIEHE
even when the growing points are not the only perceptive regions they exercise a
80
controlling influence over the resulting movements of curvature J ).

The capacity of organs to assume a definite direction by means of

curvatures of growth is distinguished, according to the nature of the
particular exciting stimulus, as heliotropism, geotropism, hy drotropism,
etc.;
and these again are either POSITIVE or NEGATIVE, according as
the direction taken by the curvature is towards or away from the
source of the stimulus. Plant organs which place themselves more
or less transversely to the line of action of the operative forces are
termed DIATROPIC. As a special result of diatropism, a transverse
position is assumed which is exactly at right angles to the direction
in which the influence which acts as the stimulus is exerted. Dorsi-
ventral organs, in particular, exhibit a tendency to assume diatropic
or transverse positions.

A. Heliotropism

The importance of light to plant life is almost incalculable. It is

not only absolutely essential for the nutrition of green plants, but it
has also a powerful effect upon the growth and general health of the
plant. Deprived of light for any length of time, leaves and
flowers usually fall oflf; fully developed, vigorous organs of green
plants soon become yellow in the dark, and droop and die. Prolonged
darkness acts like a poison upon those portions of plants accustomed
to the light. On the other hand, exactly the reverse is true of plants
or organs whose normal development is accomplished in darkness.
 PHYSIOLOGY 275

Upon them the light has a most injurious, even fatal, effect, as may
be easily observed in the case of Fungi and Bacteria. The hygienic
importance of daylight in dwelling-places is due to the destructive
action of light upon such forms of plant life. That some plants seek
the light, while others avoid it, is not surprising in view of the
adaptability which organisms usually exhibit in respect to the in-
fluences with which they come
in contact in the natural course
of their development.
Agood opportunity for the
observation of heliotropic pheno-
mena is afforded by ordinary
window-plants. The stems of
such plants do not grow erect
as in the open air, but are in-
clined towards the window, and
the leaves are all turned towards
the light as if seeking help.
The leaf-stalks and stems are
accordingly POSITIVELY HELIO-
TROPIC. In contrast with these
organs the leaf-blades take up
a position at right angles to the
rays of light in order to receive
as much light as possible. They
are diaheliotropic, or TRANS-
VERSELY HELIOTROPIC, in the
strictest sense (Fig. 223). If

among the plants there should

be one with aerial roots, Chloro-
phytum for instance, an example FIO. 2-23. A sci-aimx of the white Mustard m a
wat " r culture w* i< 1 has flrst bee illumi " :ltt"
Of NEGATIVE HELIOTROPISM will
1

?
,
. . . from all sides and then from one side only. 1'hc
be afforded, as the aerial rOOtS stem is turned towards the light, the root away
Will be found tO grow away from 'k> wlli ''' tin- leaf-blades are expandi-d ;it

from the window and turn

towards the room.
For more exact investigation of heliotropic movements it is neces-
sary to be able to control more accurately the source and direction of
the light. This can be best accomplished by placing the plants in a
room or box, lighted from only one side by means of a narrow opening
or by an artificial light. It then becomes apparent that the direction
of the incident rays of light determines the heliotropic position;
every
alteration in the direction of the rays produces a change in the posi-
tion of the heliotropic organs. The apical ends of
positively heliotropic
organs will be found to take up the same direction as that of the rays
of light.
276 BOTANY

The exactness with which this is done is illustrated l>\ an experiment made
with Pilololits criisfiflfimtx 'Fig. 224). The sporangiophorcs of this Fungus arc
quickly produced on moist horse or cow dung. They arc positively heliotropic,
and turn their black sporangia towards the source of light. When ripe these
sporangia are shot away from the plant, and \\ill be found thickly clustered about
the centre of the glass covering a small aperture through which alone the light has
been admitted a prool that the sporangiophores were all previously pointed
:

8I
exactly in that direction ( ).

Upon closer investigation of the manner in which the POSITIVE

CURVATURE of an organ is accomplished, it is found TH.VI
THE SIDE TURNED TOWARDS
THE LIGHT GROWS MORE
SLOWLY, THE SIDE AWAY
FROM THE LIGHT MORE
RAPIDLY THAN WHEN ILLU-
MINATED FROM ALL SIDES.
This may be readily shown
by previously marking with
Indian ink regular intervals
from one to two millimetre.-
apart on the opposite sides
of the organ. After the
curvature has taken pl.-ue
the intervals between the
marks will be found to it- 1

much farther apart on tin-

shaded side than on the
side turned to the light.
Fit;. L'lM. r;ii,l,nhi* :i"!i.<t<tlliini.*(I'). a nx its s]Hiiaiii:i;i As compared with the
tonanis tin; li^lit. (!, Sheet nt' ^lass /;. <i]mque case
with a circular npiMiin.L,' at 1- M. v : ssel containing elongation under normal
hiirse.(liin<;. (Cf. (leseri]ition in text.) conditions of growth, the
marks on the illuminated
side have remained nearer together, while those on the shaded side
have drawn farther apart that is, the growth in the case of a positive
;

heliotropic curvature has been retarded on the illuminated side and

promoted on the shaded side. It also becomes evident, from observa-
tion of the ink-marks, THAT CURVATURE TAKES PLACE ONLY IN THE
PORTIONS OF STEMS STILL IN PROCESS OK GROWTH. AND THAT THE
CURVATURE IS AS A RULE GREATEST WHERE THE GROWTH IS MOST
VIGOROUS.
It was formerly believed that the increased growth of the shaded side \\as pro-

duced by the beginning of etiolation, and that the diminished growth on the illumin-
ated side was due to the retarding effect which light exerts upon growth in length
(p. 258). Other heliotropic phenomena were found to be at variance with thi-.
explanation of hcliotropisin. Unicellular perfectly transparent fungal hypha- are
also subject to positive hcliotropic curvature, although in this instance there can
be no shaded side: on the contrary, the side of a hypha turned away. from the
 PHYSIOLOGY 277

light is especially illuminated on account of the refraction of the light rays. The
fact, too, that negative heliotropic curvatures also take place renders it evident that
heliotropism cannot be due to one-sided etiolation for in negative heliotropism
;

the side most directly illuminated is the one that grows more rapidly, although
tin-
retarding effect of light on the normal growth in length of negatively heliotropic
organs is
equally operative (roots, rhizomorpha).

It is evident from these considerations that it is not the difference

in the intensity of the light which causes the heliotropic curvatures,
but the direction in which the most intense light rays enter the organs.
LIGHT ACTS AS A MOTORY STIMULUS WHEN
IT PENETRATES AN ORGAN
IN ANY OTHER DIRECTION THAN THAT WHICH CORRESPONDS WITH THE
POSITION OF HELIOTROPIC EQUILIBRIUM.

The heliotropic curvatures are most strongly produced, just as in the case of
the heliotactic movements of freely moving swarm-spores, by the blue and violet
rays, while red and yellow light exerts only an extremely slight influence, or none
at all. It is due to the fact that the red-yellow and blue-violet rays are always

present together in daylight, that the heliotropism of the leaves is of advantage to

their assimilatory activity. Intermittent illumination has a more powerful effect
on the plant than light of uniform intensity.
Sensibility to heliotropic influences is prevalent throughout the vegetable
kingdom. Even organs like the roots of trees, which are never under ordinary
circumstances exposed to the light, often exhibit heliotropic irritability. Positive
heliotropism is the rule with aerial vegetative axes. Negative heliotropism is
much less frequent ;
it is observed in aerial roots, and sometimes also in climbing
roots (Ivy, Ficus stipulata, Begonia scandcns), in the hypocotyl of germinating
Mistletoe, in many, but not all, earth roots (Sinapis, ffelianfJnts), in tendrils
(chiefly in those with attaching discs), and in the stems of some climbers. By means
of their negative heliotropic character, the organs for climbing and attachment turn
from the light towards, and are pressed firmly against, their support.
Negative heliotropic curvatures are occasionally produced, not in the region of
most rapid growth, but in the older and more slowly growing portions of the stem.
The stems of Tropaeoliimmajus, for example, exhibit positive heliotropic curvatures
in the region of their greatest elongation, but lower down the stems, where their

growth is less rapid, become negatively heliotropic.

TRANSVERSE HELIOTROPISM is confined almost

solely to leaves and
leaf-like assimilatory organs, such as Fern prothallia and the thalli of
Liverworts and Algae. In these organs transverse heliotropism, in
conformity with its great utility for assimilation, predominates over
all other motory stimuli. Thus it is possible to cause the leaf-blades
of a Malva or a Tropaeolum to turn completely over
by illuminating
their under surfaces by means of a mirror.

In too bright light the transverse position of the leaves becomes changed to a
position more or less in a line with the direction of the more intense light rays.
In assuming a more perpendicular position to avoid the direct rays of the mid-day
sun, the leaf-blades of Lactuca Scariolu and the North American Silphium h/r /'///'>/-
turn necessarily take, according to STAHL, the direction of north and south, and so
278 1JOTANY PART i

arc often referred to as COMPASS PLANTS. A vertical jtosition is also attained hy

development of phyllodes, in connection with which may 1.x- mentioned the
tli<'

82
vtTtically placed leaves of many Myrtaceae and Proteaceae ( ).

The heliotropic character of organs may change through the

activity of external influences, and also at different stages of their
development and growth. The flower-stalks of JJnin-in cymbaiaria
are at first After pollination, however, they
positively heliotropic.
become negatively heliotropic, and as they elongate they push their
fruits into the crevices of the walls and rocks on which the plant
(p. 257). The intensity
^rnws of the illumination has a great influence,
since plants which in subdued light are positively heliotropic exhibit
negative heliotropism when the illumination is excessive. Between
the two reactions a neutral aheliotropic condition exists. OLTMANNS.
who elucidated this variation of the reaction with the intensity of
the light, and the search for an optimal intensity which is connected
with it (cf. heliotactic swarm-spores, p. 265), termed it PHOTOMETRY.
It remains to be noted that it is not so much the absolute intensity
of the light which is of importance, but that the degree of change in
light intensity needed to alter the reaction of the plant depends on
the illumination to which the latter has been previously exposed ( 83 ).
Heliotropic sensibility is markedly increased when traces of coal-
gas, etc. are present as an impurity in the atmosphere. This is so
strikingly the case that conclusions as to' the degree of impurity can
be drawn from the heliotropic deflection exhibited by susceptible
plants (Peas, Vicia calcarata, etc.).

B. Geotropism

That the stems of trees and other plants should grow upwards and
their roots downwards, is such a familiar occurrence and so necessary
for the performance of their respective functions as to seem almost a
matter of course. Just as in the discovery of gravitation 240 years
ago, it required an especially keen spirit of inquiry to lead to the in-
vestigation of this everyday phenomenon. The fact that everywhere
on the earth, stems take a perpendicular direction and that, while
;

buried in the earth, this same direction is assumed with certainty by

germinating seeds and growing shoots and chiefly the fact also that
;

a shoot, when forced out of upright position, curves energetically

its
until it is again perpendicular, led to the supposition that the cause
of these phenomena must be in a directive force proceeding from the
"
earth itself. The correspondence in the behaviour of a stem in
always assuming a perpendicular position, with the continued
maintenance of the same direction by a plumb-line, suggested at once
the force of gravitation, and the English investigator KNIGHT, in
1809, demonstrated that the attraction of gravitation, in fact, exerted
an influence upon the direction of growth. As KNIGHT was not able
SECT, ii PHYSIOLOGY 279

to nullify the constantly operative influence of gravity upon plants

and so directly prove its influence, he submitted them to the action
of centrifugal force an accelerative force operating like gravity
upon the masses of bodies, which had, in addition, the advantage
that it could be increased or diminished at will. KNIGHT made use
of rapidly rotating, vertical wheels, upon which he fastened plants
and germinating seeds in various positions. The result of his
experiments was that the stems all turned towards the centre of the
wheel and the roots directly away from it. On wheels rotating in a
horizontal plane, where, in addition to the centrifugal force, the one-
sided action of gravitation was also still operative, the shoots and
roots took up an intermediate position depending on the combined
influence of the two forces. In this way it was positively ascertained
that terrestrial gravitation determines the positions of plant organs
in respect to the earth ( 84 ).
Later, it was also shown that not only the perpendicular direction
of stems and primary roots, but also the oblique or horizontal direction
taken by lateral branches, lateral roots, and rhizomes, is due to a
peculiar reaction towards the force of gravitation.
The property of plants to assume a definite position with respect
to the direction of gravitation is termed GEOTROPISM. It is customary
also, as inthe case of heliotropism, to speak of positive and negative
geotropism, diageotropism,> and transverse geotropism, according to
the position assumed by the plant or organ with respect to the centre
of the earth. Still another form of geotropic irritability, lateral

geotropism, renders possible the twining of stem-climbers.

Negative Geotropism. All vertically upward growing organs, whether stems,
leaves (Liliiflorae), flower-stalks, parts of Howers, or roots (such as the respiratory
roots of Avicennia (Fig. 213), Palms, etc.), are negatively geotropic. In case
such negatively geotropic organs are forced out of their upright position, they
assume it again if still capable of growth. As in heliotropism, GEOTKOPIC CURVA-
TURE RESULTS FROM THE INCREASED GROWTH OF ONE SIDE AND THE RETARDED
GROWTH OF THE OPPOSITE SIDE, and the region of greatest growth is, in general,
also that of the greatest curvature. In negatively geotropic organs, growth is
accelerated on the side towards the earth on the upper side it is retarded. In
;

consequence of the unequal growth thus induced, the erection of the free-growing
extremity is effected. After the upright position is again attained, the one-sided
growth ceases and the organ continues to grow in an upward direction.
The process of negative geotropic movement is dependent (1) upon the vigour
:

of the existing growth ; (2) upon the sensibility of the organ ; (3) upon the fact
that the stimulus of gravity works most energetically when the apex of the ortho-
tropic organ is removed about 90' from the vertical ; the more nearly the zone
capable of curvature approaches this position, the stronger is the motory stimulus ;

deviations from this optimal position of stimulation do not result in such striking
differences in reaction as are causedby even very small (|) deviations from the
vertical ; (4) and also upon the fact that after a stimulus has ceased to act
upon a
plant, the induced stimulation continues to produce so-called AFTER EFFECTS, just
as by a momentary stimulus of light an after-perception persists in the eye.
280 BOTANY

These considerations determine tin- actual

course of the directive movement of geotropum,
which, as will be seen from the adjoining figure
(Fig. 225), does not consist merely of a simple.
continuous curvature. The numbers 1-16 show,
diagrammatically, different stages in the geotropic
erection of a seedling growing in semi-darkness
and placed in a horizontal position (No. 1). The
growth in the stem of the seedling is strongest.
just below the cotyledons, and gradually decreases
towards the base. The curvature begins accord-
ingly close to the cotyledons, and proceeds grad-
ually down the stem until it reaches the lo\\ei.
no longer elongating, jxirtions. Through the
downward movement of the curvature, and partly
also through the after-effect of the original

stimulus, the apical extremity becomes bent out

of the perpendicular (Xo. 7 and in this way
1

a curvature in the opposite direction takes place.

Thus, under the influence of the stimulus, the
stem bends back wards and forwards, until, iinally,
the whole growing portion becomes erect and no
longer subject to the one-sided action of the
geotropic stimulus.
Positive Geotropism is exhibited in tap-roots,
in many and in the leaf-sheaths of
aerial roots,
the cotyledons of many Monocotyledons which
penetrate the earth during germination. All
these organs, when placed in any other position,
assume a straight downward direction and after-
wards maintain it. Formerly, it was believed
that this resulted solely from their weight and
the pliancy of their tissues. It is now known

that this is not the case, and that positively

geotropic, like negatively geotropic.
movements
are possible only through growth. The power
of a downward curving root-tip to penetrate
mercury (specifically much the heavier... and to
overcome the resistant pressure, much greater
than its own weight, proves conclusively that
is a manifestation of a vital
positive heliotropism
process. Positive geotropic curvature is due to
THK GUOWTH OK AN UI:<;AN IN
the fact that
Kio. '2-2i>. Different stages in the pro. LENGTH <>N THK VITKI:
IS PKOMiilKl' SIDE,
i I-NS of ^eotropic movement. The AND RETARDED, T-\ IN' MnliK. STRONGLY. ON THK

SIDE TURNED TOWARDS THE EARTH. A young

figures, 1-16, indicate successive
stages in the geotropic curvature of
r, K rown in sen.i-darkness:
llinK
root of the Chestnut, growing vertically, elon-
20 mm.; when
at 1, placed horizontally; at 16, ver- gated equally on all sides
tical. For description of intermedi-
horizontally, it exhibited a growth of 28
placed
ate stages see text. (Diagrammatic.)
mm OQ the an(1 of on]y 9
upper mm
oll t lic .

lower side. Fig. 226 illustrates the way in which the curvature takes place in the
 PHYSIOLOGY 281

i-asr of a root of the Broad

Bean, which was marked at regular intervals with Indian
ink and placed horizontally.
Diageotropism. Most lateral branches and roots of the first order are diageo-
tropic, while branches and roots of a higher order stand out from their parent organ
in all directions. DIAGEOTKOPIC ORGANS ARE ONLY IN A POSITION OF F.^UI-
I.IIIRIt'M WHEN THEIR LONGITUDINAL AXES FORM A

DEFINITE ANGLE WITH THE LINE OF THE ACTION OF

GRAVITY. If forced from their normal inclination they
return to it by curving. A special instance of diageo-
tropism is exhibited by strictly horizontal organs, such
as rhizomes and stolons, which, once they have attained
their proper depth, show a strictly TRANSVERSE GEO-
TROPISM, and, if removed from their normal position,
their growing tips always return to the horizontal.
The oblique position naturally assumed by many organs W
is in part the result of other influences
(epinasty and
hyponasty, their own weight, and other directive forces).
A more complex form of geotropic orientation is
manifested by dorsiventral organs, e.y. foliage leaves,
xygomorphic flowers. All such dorsiventral organs,
just as radial organs that are diageotropic, form a definite
angle with the direction of gravity, but arc only in
equilibrium when -the dorsal side is uppermost. If, in
Flli
spite of the proper inclination of the longitudinal axis, ;
ture of the root < it ;i
seedling
the dorsal side should he underneath, the latter elon- of Vma Fl<. I, Placed
gates until it comes back again into a dorsal position. liorixnntally : II- after
seven hours ; ///. after

A state of torsion often results from the twenty-three hours

fixed index. (After SA.
; /. .1

!!*.)
orientation movements of dorsiventral organs
to recover from abnormal positions. Similarly, a torsion must also,
of necessity, occur when a geotropic organ, which has become curved
over toward its parent axis, turns itself about so as to face outwards
(EXOTROPISM) (Fig. 227).

The rotation of the ovaries of many Orchidaceae, of the flowers of the

Lobeliaceae, of the leaf-stalks on all hanging or oblique branches, of the origin-
ally reversed leaves (with the palisade parenchyma on the under side) of the
Alstroemeriae, and of Allium ursinum, all atford familiar examples of torsion
85
regularly occurring in the process of orientation ( ).

Twining Plants. In addition to the better-known forms of

geotropism already stem -climbers exhibit a peculiar
mentioned,
geotropic movement, by means of which they are enabled to twine
about upright supports. This movement depends upon the geotropic
promotion of the growth of one side (not, as in negative or positive
geotropism, of the upper or lower portions). Thus a geotropic
curvature in a horizontal plane is produced (LATERAL GEOTROPISM),
resulting in a revolving motion of the shoot apex. Twining plants
occur in very different plant families and although an upward
;

growth is essential to their full development, their stems are not able
282 BOTANY PART I

of themselves to maintain mi erect position. The erect stems of other

plants, which often secure their own rigidity only through great

expenditure of assimilated material (in xylem and sclerenchyma) are

made use of by stem-climbers as supports, on which to spread out their
assimilatory organs in the free air and light. The utilisation of a
support produced by the assimilatory activity of other plants is a
peculiarity they possess in common with other climbers, such as
tendril- and root-climbers. Unlike them, however, the stem-
climhers accomplish their purpose, not through the use of lateral
clinging organs, but by the capacity of their main stems to twine
about a support. The first internodes of young stem-climbers, as

Ki;. -_7. The movements by which alflow-r <it Ai-imittnit iiii/llns nonius its proi-i po.Mtion when
the axis bearing it (S)is inverted. I, Inverted position II, position resulting from gi-otrnpisni.
;

the flower facing the parent axis; ///, flower .again facing outwards. aftT thf exotropic
movement.

a rule, stand erect. By further growth the free end curves

energetically to one side, and assumes a diageotropic, more or less
oblique or horizontal position. At the same time the inclined apex
begins to revolve in a circle either to the right or to the left. This
is the movement which it has been customary to speak of as
"
revolving nutation," but which it is better to term REVOLVING
MOVEMENT. The expression " nutation " is not in this case correct, as
by it are understood autonomic movements ; while THE REVOLVING
MOVEMENTS OF STEM-CLIMBERS RESULT FROM THE EXTERNAL
STIMULUS OF GEOTROPISM, which causes a promotion of growth in
either the right or left side of the young internodes of the inclined
shoot apex. As a result of this, a movement towards the other side
is induced. On account of the direct connection of the apex of the
shoot with the lower erect internodes, this revolving movement
necessarily gives rise to a similar rotation of the revolving apex on its
 PHYSIOLOGY 283

longitudinal axis. Through this rotation the torsion, which would

otherwise be produced by the revolving movement of the inclined tip
of the shoot, is released. (This process will at once become apparent
by imitating the movement with a rubber tube.) Thus the apex of
a stem-climber sweeps round in a circle like the hands of a watch,
and rotates at the same time like the axle to which the hand is
attached. By this rotation of the shoot apex, the part of the
stem subjected to the action of the lateral geotropism is constantly
changing ; and the revolving movement once begun, must continue, as
no position of equilibrium can be attained.

Without the constant and unchanging action of gravitation in determining the

direction of the revolving movement, the twining of a shoot continuously about a
support is hardly conceivable. It is accordingly not without reason that the

revolving movement is a continuous, fixed, geotropic movement, and not an

autonomic nutation without definite directive force. Lateral geotropism is a
physiological requisite for the climbing, and the existence of stem-climbers as such
isdependent upon this peculiar form of geotropism. To this dependence, however, is
also due the fact that stem-climbers can only twine about upright or slightly
inclined supports. This is, it is true, a limitation to their power of climbing, but
one which is not without advantage, for the plants are thus constrained to ascend
to freer light and air ( S6 ).
When an upright support occurs anywhere in the immediate neighbourhood of
the apex of a climbing shoot it is sure to be discovered. The apical extremity, the
movement of which is but little disturbed by the leaves, which remain for a long
time undeveloped, is forced by its lateral geotropism against the support, and by
itsnext revolutions twines around it. If the support be thin, the coils, at first
almost horizontal, are only loosely wound about it. Later they become more
spiral,and so wind more tightly. This is accomplished by the ultimate predomi-
nance of negative geotropism in the coiled portions of the stem, which tends
continually to draw out the coils and make the stems upright. This action of
negative geotropism is well shown in the case of shoots which have formed free
coils without a support. By the resistance offered by the supports to the
complete elongation of the spiral stems, the shoots are held firmly in position. In
many twining plants the roughness of their surfaces (due to hairs, bristles, hooks,
furrows) also assists in preventing the shoots sliding down their supports. The
autonomic torsion arising in the older portions of the stems is also of assistance in
holding climbing plants, especially those with furrowed stems, tightly wound about
their supports. The twining of stem-climbers, as well as the attachment to their
supports, is thus due to geotropic processes of growth, and not, as in tendril-
climbers, to contact stimuli. By the mechanism of their climbing process twining
plants are restricted to moderately stout supports.
In addition to the autonomic torsions, a torsion from purely mechanical causes
is necessarily manifested in the elongation of the coils of a twining stem, which are

at first nearly horizontal, so far at least as it is not equalised by the free movement
of the apex.
The direction of the revolving movements, and accordingly also of the windings,
of most stem - climbers is constant. The twining stems are for the most part
SIXISTHORSE (Convolvulus, Phaseolus, P/tarbitis, etc.). Seen from above, the wind-
ings run from the north towards the west, south and east to north again, i.e. just the
284

rever.-e of the movement of the hands of ;i watch. Viewed from the side, the wind-
ings ascend the support from the left below to the right above (Fig. 228). Di.x-
TKOKSE stem-climbers .with windings from east to west occur less fre<[iiently (Hop,
Honeysuckle, Polii<j<nini<i cmu-ofm/iis, etc.). In the example chosen for illustrati<m
(Mi/r.-i/p/ii/lliim <isi>n,-ij<>i<lrs, Fig. 229) the undeveloped condition of the lateral
members in comparison with the elongated internodes of the stem is very apparent.
A very few plants, such as Blwntnbtuhia lateritia, Jlil/lii /// i/i/i/n/'i, and Sci/~
si-cm able to climb equally well either to the right or to the left.
. A

Ik

Fi<;. -22s. A siniatrorM stem-cumber, .. _'_".'. A dextrorse stcin-c-liiiilii-i-.

I'lmrbiti* lii*jiiila. Tin- upper Myi-xilihyllii'iii ti&paranniilr-<. Tin-

leaves remain small for a lon^ short lateral slioots liavi- de-
veloped phyllocladia.

similar irregularity is shown in Solanuai dulcamara, \\liich, however, rarely

twines, and then only under special circumstances.
When the apex of a sinistrorse shoot points towards the north, it is the growth
of the east side which is promoted by geotropism in dextrorse climbers, on the
;

contrary, the growth of the west side is more rapid. From the fact that the
promotion of growth occurs always on the same side, it will be apparent that the
8T
apex of an inverted twining stem must unwind from its support ( ). (Concerning
the behaviour of stem-climbers on the klinostat compare p. 287).
Curvature of Grass - Haulms. All the examples of geotropic movements, so
 PHYSIOLOGY 28;

far observed, took place ill the growing portions of plants, and were due to a
disturbance of the course of growth. A curvature even of lignified twigs can also
be produced by the one-sided stronger growth of the cambium and of the young
secondary tissues. Even many -year-old branches of Conifers are still able, although
exhibit geotropic curvatures ( 8S ).
.slowly, to THE NODES OF GRASSES SHOW THAT
BESTING TISSUES ALSO CAN HE EXCITED TO NEW GROWTH BY THE STIMULUS OF
GRAVITATION. The knot-like swellings on the haulms of the Grasses are not nodes
in a morphological sense, but are barrel-shaped thickenings of the leaf-sheaths above
their actual insertion on the shoot axis. The part of the stem thus enveloped is
very tender and flexible. When a grass-haulm is laid horizontally, which not
unt'requently occurs through the action of the wind or rain, the nodes will begin
to exhibit an energetic growth on their lower sides. As the upper sides of the
nodes take no part in the growth, but are instead frequently shortened through

Km. _>:>. (ieotropic erection n( a grass-haulm

liy tin- curvature of a node. 1, Placed
horizontally, lx.it h sides (. ./) of the node

being of equal length ; '2, the under side

() lengthened, the upper side (o) somewhat Fli:. _':!!.- Seedling of ' '< rhi/n. The peg
shortened as a result of the curvature
;
(W) (levelo)ied from the under side of the
the grass-haulm has been raised thrmigli hyjiocotyl assists in rupturing the seed-
an angle of 75. coat.

pressure, and lo>s (if water, knee-like curvatures are formed at the nodes, by means
of which the haulm isagain quickly brought into an erect position (Fig. 230).
Seedlings of Cucurbitaceae. The stimulus of gravity induces the growth of a
peculiar peg-like outgrowth from the lower side of the hypocotyl of the seedling of
<.'iu-in-lnta (Fig. 2:31 W
This peg, which results from a limited
.

geotropic growth
in thickness, assists in liberating the cotyledons from the seed-coat.

Modifications in the character of the geotropism, as of the helio-

tropism, of an organ may be occasioned by the operation of internal
as well as external influences. Such changes in their geotropic
position frequently occur, as VOCHTING has demonstrated, during the
development of flower-buds, flowers, and fruits, whose movements
are,according to WIESNER, especially influenced by epinastic load-
curvatures (p. 272). The burial of the fruits of Trifolium suUerr,i<in,<
and Anichis Itj/iiogaea is due to geotropism. Of the
changes in the
geotropic conditions of plant organs due to external causes, those are
particularly noticeable which result from a failure of a sufficient
supply of oxygen, by which, for example, roots and rhizomes are
made negatively geotropic. The temperature influences the position
286 BOTANY PAKt i

of the main or lateral axes of many plants. Thus the stems of many
when the temperature soil and
is low, lie flat on the
spring plants,
only assume the erect position as the result of geotropism when lie t

temperature becomes higher. Even more important are the modi-

fications arising from the action of light, by which the geotropir

irritability of rhizomes and foliage leaves may be so modified or

weakened as to permit of more advantageous heliotropic positions.
While heliotropism was increased by the impurity of the atmo-
sphere (p. 278), geotropism is weakened by this influence.

C. Hydrotropisin, TJii'i'iitii/r<>j>isin.
n n<l nil,, / 7V( //*/.>;//.<

While the illumination and the position with regard to the earth are the most
general and important relations of the plant to its environment, and heliotropism
and geotropism the most widely spread reactions of the plant, they are not tin- only
phenomena of the kind. Whenever any external force or substance is important
to the vital activity of a plant or any of its organs, there will also be found to be

developed a corresponding irritability to their influences. Roots in dry soil are

diverted to more favourable positions by the presence of greater quantities of
moisture. The force of this POSITIVE HYDROTROPISM may be so great as to over-
come the geotropic equilibrium of the roots, and thus give rise to hydrotropic
curvatures. Conversely, the sporophores of many mould Fungi avoid moisture.
To this property is due the fact, so advantageous for the distribution of the
spores, that their sporangiophores grow directly away from a moist substratum.
Corresponding to the chemotactic irritability of Bacteria and spermatozoids, roots,
fungal hyphse, and pollen tubes exhibit positive and negative CHEMOTIUHMC
CURVATURES. These vary according to the concentration of the solution, .so that
an attractive substance, at a higher concentration, may act repulsively. THERMO-
TROPISM or caloritropism (due to the stimulus of heat), RHEOTROPISM (occasioned
by the direction of water currents), and AEROTROPISM (a form of chemotropism)
are additional phenomena, which have been distinguished as arising from the
special action of external stimuli, and stand in direct relations to certain vital
89
requirements of plants ( ).
In the case of EI.EI.TROTROPISM, which has also been demonstrated in plants, i,<>
such essential relations have been discovered; the disposition of plant organs in
a direction contrary to that of an electric current seems in no way to affect th oil-
well-being. The fact of the existence of electro tropism in plants shows clearly that
an irritability may be present, from which no direct benefit is ordinarily derived,
and which accordingly could not have been attained by natural selection.

I). The Method of Slow Rotation The Klinostut

All the curvatures of growth previously discussed have been induced by the
one-sided action of stimuli, the source of which determined the direction of the
movements as well as the position of equilibrium. An influence operating eqiiallv
on unable to produce a curvature in an organ of which the irritability
all sides is
is equally developed on all sides. In like manner no curvatures can take place
when the plant is uniformly rotated, with a velocity sufficient to preclude the
continuous operation of a stimulus on any one point long enough to occasion a
one-sided growth. As in that case, no one side will be exclusively acted upon,
SECT, ii PHYSIOLOGY 287

but the growth of all will he equally promoted or retarded, the action of external
influences, although exerted in only one direction, will be equalised. On this
account the "method of slow rotation," originally instituted by SACHS, is of great
assistance in the observation and investigation of the phenomena of movements.
By means of it, heliotropic movements due to one-sided illumination may he
prevented, without the necessity for either exposing the plants to the injurious
effects of continued darkness or providing for an equal illumination on all sides.
This method is, moreover, of especial value in investigating the movements due to
the action of gravitation, for it is not possible to exclude its influence, as it is

those arising from light, definite temperature, oxygen, etc.

WHEN PLANTS ARE SLOWLY ROTATED ON A HORIZONTAL AXIS THE ONE SIDED
ACTION OF GRAVITATION IS ELIMINATED, AND GEOTROPIO CURVATURE IS THUS
PREVENTED in organs which react equally on all sides. The rotations are best
produced by the KLINOSTAT, an instrument by means of which an exactly horizontal
axis is rotated by clock-work. That geotropic curvatures of radial organs are, in
fact, precluded by means of the klinostat, furnishes a remarkable corroboration
of the result of KNIGHT'S experiments, and is a further proof that such
-curvatures are due to terrestrial gravitation. Through the equalisation of
the action of external directive influences, radial portions of plants exhibit on
the klinostat only such movements as arise from internal causes. The most
important of these autouomic movements are those resulting in epinastic and
hyponastic curvatures (p. 272), and the retrogression of recently formed paratonic
curvatures through longitudinal extension (autotropism) or by curvature in the
90
opposite direction ( ).
Such autonomic movements should not be confused witli those exhibited by
dorsiventral organs on the klinostat, in consequence of the unequal irritability of
their different sides. Through the special irritability of the dorsal side (p. 281)
of foliage leaves and zygomorphic flowers, it is during their rotation more strongly
acted upon by geotropic influence than the ventral side ; as a result of this, curva-
tures are produced which so closely resemble those resulting from epiuasty that
they were for a long time actually considered as such ( ). When stem-climbers
91

are rotated on the klinostat, their revolving movement ceases, the part of the stem

capable of growth unwinds and straightens, and afterwards exhibits only irregular
92
nutations ( ).

E. Curvatures induced by Contact Stimuli

The protoplasm of plants, like that of animals, exhibits an irrit-

ability to contact, whether momentary or continuous. This is

apparent, not only from the behaviour of naked protoplasmic bodies,
but also from the reactions manifested by walled cells and by whole
organs, the functions of which may be so disturbed by the action of
mechanical stimuli that death ensues. The almost universal irritability
of vegetable protoplasm to mechanical stimulation is utilised by a
number of plants for the production of movements which lead to their
ultimate attachment to the irritating body. The mechanical stimulus
is frequently perceived by means of special sense organs, the protoplasts
of which are altered in shape owing to the contact. Tendril-climbers,
in particular, have developed this irritability to contact stimuli as a
means of attaching themselves to supports (cf. p. 67, Fig. 69); and
288 BOTANY PAKT i

in that way are enabled to elevate their assimilating and also their
reproductive organs into more favourable situations. In the case of
twining plants which possess similar powers of climbing, the process
of elevation, as has already been shown, is accomplished by means of
the geotropic irritability of the stems themselves. In the case of
tendril-climbers, on the other hand, the attachment to the support is
effected, not by the main axis of the plant, but by lateral organs of
different morphological character. These may either maintain, at
the same time, their normal character and functions, or, as is usually
the case, become modified and as typical tendrils serve solely as
climbing organs. According to FITTING, contact with a solid body
quickly induces an increase in the growth of the opposite side of the
organ, and this without any retardation of growth on the touched
side leads to a sharp curvature of the tendril which coils it
about the support. The more slender the tendrils and the stronger
their growth, the more easily and quickly this process occurs.

Through the tendency of the curvature to press the tendrils more

and more firmly against the support, deep impressions are often made
by them upon yielding bodies, soft stems, etc.
According to PFEFFER'S investigations, it is of great importance to
the tendrils in the performance of their functions that they are not
induced to coil by every touch, but only through CONTACT WITH THE
UNKVEN SURFACE OF SOLID BODIES. Rain-drops consequently never
act as a contact stimulus ; and even the shock of a continued fall
of mercury produces no stimulation, while a fibre of cotton-wool

weighing 0'00025 mgr. is sufficient to stimulate the tendril ( ).

9:!

In the more typically developed tendrils tlic curvature does not remain restricted
to the portions directly subjected to the action of the contact stimulus. Apart
from the fact that, in the act of coiling, new portions of the tendrils are being
continually brought into contact with the support and so acted upon by the
stimulus, the stimulation to curvature is also transferred to the portions of the
tendril not in contact with the support. Through the action of the propagated
.stimulus, not only is the free apex of the tendril turned more quickly around the
support, but a tendency to curvature is imparted to the portion of the tendril
between the support and the parent shoot. As this is extended between two fixed
jxiints, tliis tendency causes it to coil spirally, like a corkscrew. With the spiral
coiling, a torsion is produced, and since, on account of the fixed position of the
two end points, it cannot be exerted in one direction only, the spiral, for purely
mechanical reasons, coils partly to the left and partly to the right. POINTS OF
UEVKKSAL (x) thus occur in the windings which, in equal numbers to the right
and to the left, equalise the torsion (Fig. 232). By the spiral coiling of tin-
tendrils the parent-stem is not only drawn closer to the support, but the tendrils
themselves acquire greater elasticity and are enabled to withstand the injurious
effects of a sudden shock.

Advantageous changes also take place in the anatomical structure of the tendrils
afterthey are fastened to the supports. The young tendrils, during their rapid
elongation, exhibit active nutations, and thus the probability of their finding a
8CET. 11 PHYSIOLOGY 289

support is enhanced. During this time they remain soft and flexible, while the
turgor rigidity of their apices is maintained only by collenchyma. In this con-
dition they are easily ruptured, and have but little sustaining capacity. As soon,
however, as a support is grasped, the coiled-up portion of the tendril thickens
and hardens, while the other part lignifies, and becomes so strengthened by
sclerenchymatous formations that the tendril can finally sustain a strain of many
pounds. When the tendrils do not find a support they usually dry up and fall off,
but in some cases they first coil themselves into a spiral.
The tendrils of many plants (Cobaea, Eccremocarpus, Cissus) are irritable and
capable of curving on all sides others (tendrils of Cucurbitaceae and others with
;

Flo. 232. Portion of a stem of S-icytm angvlutua with

tendril ; x, point of reversal in the coiling of the tendril.

hooked tips) are, according to FITTING, sensitive

on all sides but only curve when the under side
is touched if the upper' surface is at the same
;

time stimulated curvature is arrested. In some FHJ. 233. Portion of a cl

the shoot of A mpdopsis Veitch ii. The

cases tendrils quickly grasp the support
tendrils (R) have fastened them-
(Passiflora, Sicyos, Bryonia) ; while in other
selves to a smooth wall by nn-ans
tendrils the supports are very slowly grasped of holdfasts.
(Smilax, Vitis).
Tendril-climbers are not, like twining plants, restricted to nearly vertical
supports, although, on account of the mariner in which the tendrils coil, they can
grasp only slender supports. A few tendril-climbers are even able to attach them-
selves to smooth walls. Their tendrils are then negatively heliotropic, and
provided at their apices with small cushion-like outgrowths, which may either
develop independently on the young tendrils, or are first called forth by contact
irritation. Through their sticky excretions these cushions become fastened to the
wall and then grow into disc-like suckers, the cells of which come into such close
contact with the supporting wall that it is easier to break the lignified tendrils
than to separate the hold-fasts from the wall. Fig. 233 represents the tendrils of
Quiiiaria (Ampelopsis) Veitchii. The suckers occur on its young tendrils in the
form of knobs. In Quinaria quinquefolia and Qu. radicantissima the suckers are
U
290 HOTANY PART i

only produced as the result of contact, and the tendrils of these plants are able
also to grasp thin supports.
Sometimes, as in the case of Lophospermum scundeiis (Fig. 234), the leaf-stalks,
although bearing normal leaf- blades, become irritable to contact stimuli and
function as tendrils. Of leaf-stalks which thus act as tendrils, good examples are
alforded by Tropaeolum, Maurandia, Solanum jasminoides, Nepenthes, etc. The
subsequent modifications occurring in more perfectly developed tendrils are not
noticeable in the case of petiolar tendrils, although the coiling portion of the leaf-
stalk of Solanum jasminoides does become strongly thickened and lignified ;

while the leaf-blades of Clematis, by remaining small for a time, enhance the
tendril-like character of their leaf-stalks,and by bending backwards also assist in
maintaining the initial contact with a support. At other times the midribs of
the leaf-blades themselves become prolonged, and assume the function of tendrils
(Gloriosa, Littonia, Flagellaria). In many species of Fumaria and Corydalis, in
addition to the leaf-stalks, even the leaf-blades of the leaflets twine around slender
supports, while the parasitic shoots of Cuscuta (Fig. 186) are adapted for both
twining and climbing. Climbing parts of the thallus occur in some Thallophyta
(Florideae) (
w ).

Kic. 1:14.- Loplioxpernnim scandens climbing by means of its tendril-lik** ]'tioles.

F. Curvatures of Growth due to Variations in Light and Temperature

The flowers and foliage leaves of many plants exhibit the

peculiarity that their different sides (the upper and under sides of
foliage-leaves and leaf-stalks, the inner and outer sides of floral leaves)
show an unequal growth in response to even transitory and slight
variations in temperature and in the intensity of light. Whenever,
on account of such variations, the growth of the under side of a leaf
exceeds that of the upper side, the whole leaf moves upwards and
 PHYSIOLOGY 291

towards the parent axis ;

while if the growth of the upper side is the
stronger, the leaf is
depressed.

Movements of this nature are especially noticeable iu flower-leaves, and bring

about the opening and closing of the flower. A rise of temperature causes the
flowers of the Tulip and Crocus, and also those of Adonis, Ornithogalum, and
Colchicum, to open, while sudden cooling causes them to close. Tulips and
Crocuses, if brought, while still closed, into a warm room, open in a very short
time with a difference of temperature of 15-20 C., in from two to four minutes.
;

Crocuses respond to an alteration in temperature of C. Tulips to a variation of ;

2-3 C. In warm' sunshine the spring or summer flowers are open for the visits of
insects, but on a lowering of temperature the sexual organs are covered up and
protected.
The flower-heads of Taraxacum, Leontodon, and other Composites, also the
flowers of Nyuiphaca, Cacti, etc.,

open when illuminated, and close

when kept in darkness (Fig. 235).
Variation of light produces also
unequal growth in foliage-leaves,
particularly in those of the Chenopo-
diaceae, Caryophyllaceae, and Balsa-
minaceae, and cause them to assume
so-called SLEEP POSITIONS.
In many instances the move-
ments of the floral leaves are pro-
duced by variations of light as well
as of temperature ;
for example, FIG. 236. Flower-head of Leontodon closed
hastilis,
some of the flowers mentioned open when kept in darkness, open when illuminated.
in the light and close in the dark, al- (From DETMER'S PhysM. Pract.)
though the temperature remains con-
stant. In the case of opposing external influences, the resulting direction of the
movement of the flower-leaves is determined by the influence which is predominant.
The dependence of these movements upon different, and often opposing, influences,
together with the continuance of movements induced by previously operative
influences, led to their being for long misunderstood, especially as the movement
of opening is from internal causes followed by a closing movement even if a high
95
temperature is maintained ( ).

These movements, occasioned by variations in the illumination

and temperature, must not be confused with those of heliotropism
and thermotropism in both of which the movement induced in an
;

organ is dependent upon its relative position with respect to the

source of the light or heat, and not upon the varying
intensity of
the stimulus. The movements last discussed are photonastic or
thermonastic, i.e., they result from the variation in illumination or
temperature, and not from the direction of the light or heat rays.

3. Movements due to Changes of Turgor (Variation Movements)

The various movements hitherto considered are, to a large extent,
292 BOTANY PART I

the result of the action of forces acting on growth. These move-

ments were therefore confined to organs, or parts of organs, still in a
state of growth. In contrast to the almost universal immobility of
all fully-grown organs, it is interesting to find that some plants have
found a means of carrying on vigorous movements without the assist-
ance of growth.
It has already been shown (p. 179) that through the pressure of

increasing turgidity the elastic cell walls become greatly distended

and the cell cavity expanded, while, on the other hand, the cell
walls shrink and the cell becomes smaller when the turgor is
diminished (Fig. 174). It is to these changes in volume, which thus
result from alterations in turgor, that the varying movements of
fully developed living organs are due.
Such variation movements occur only in foliar organs (foliage
and flower leaves, stamens, style and stigmatic lobes). These move-
ments are especially noticeable in the compound leaves of the
Leguminosae and Oxalideae, and also in the leaflets of Mtirsilui. In
the motile regions of these leaves special masses of tissue are, both
physiologically and anatomically, adapted for the promotion of this
form of movement.

This tissue appears externally as a firm cushion or PULVINUS, sharply distin-

guished from the rest of the leaf-stalk, and is the direct cause of the leaf move-
ments. Anatomically considered, the pulvinus consists, for the most part, of
strongly turgescent parenchyma with very elastic cell walls. The vascular bundles
and mechanical elements, which, in other portions of the leaf-stalk, are situ-
ated towards the circumference, unite in the pulvinus in the form of a single flex-
ible strand, and so offer little opposition to the movement of the leaf resulting from
the curvature of the motile region (p. 182). The unfavourable arrangement of the
mechanical tissues is compensated by the turgescence of the pareuchymatous
tissues 6n which the support of the leaf-blade in its proper position in these cases

depends. The parenchyma of the pulvinus forms a thick enveloping layer about
this axial strand, by means of which, through the pressure arising from a differ-
ence in the turgescence of its opposite sides, a movement of the whole leaf-blade
is brought about, similar to that of the outspread hand by the motion of the wrist.

These variation movements are either autonomic, when the varia-

tions of turgor are due to no recognisable external influence, or
paratonic (aitionomic), when the turgor is regulated in a definite
way by the action of external stimuli.
Autonomie Variation Movements. A remarkable example of
this form of movement is furnished by the small lateral leaflets of
Desmodium (Hedysarum) gyrans, a papilionaceous plant growing in the
damp Ganges plains and still more strikingly, according to
MOLISCH, by Oxalis hedysaroides. In a moist, warm atmosphere (22-
25) these leaflets make circling movements which are in no way
disturbed by variations in the intensity of the light, and are of such
rapidity that the tips describe a complete circle in 1-3 minutes; in
 PHYSIOLOGY 293

the case of the leaflets of Omlis a path of |-li cm. is traversed in one
or a few seconds. Variations in illumination do not disturb these
movements. The autonomic variation movements of Trifolium and of
the Wood-Sorrel take place, on the contrary, only in darkness. Thus
the terminal leaflets of Trifolium pratense exhibit oscillatory movements
in the dark with an amplitude which may exceed 120, and are
regularly repeated in periods of 2-4 hours but on exposure to light
;

the leaflets cease their oscillations and assume a fixed light position.
Paratonie Variation Movements are chiefly induced by variation
in the intensity of the light, by the stimulus of gravitation, and by
mechanical irritation (shock, The pulvini of leaves may be
friction).
affected by several the leaves of Mimosa pudica, for
different stimuli ;

example, are set in motion by the action of light, by the stimulus of

a shock, and sometimes also by gravitational stimuli ; in addition
they exhibit autonomic movements.
A change from light to darkness, as from day to night, occasions
NYCTITROPIC MOVEMENTS or the so-called SLEEP MOVEMENTS. In the
day or light position, which is the same as that of diaheliotropic
foliage-leaves, the leaf-blades are perpendicular to the incident rays
of light. With the commencement of darkness the leaves or the
single leaflets fold either upwards with their upper surfaces inward,
or downwards with their lower surfaces together, and so remain until
the diurnal position is again assumed on recurring illumination.
THE VARIATIONS IN TURGESCENCE IN THE OPPOSED HALVES OF THE
JOINT STAND IN INVERSE RELATION TO ONE ANOTHER OR IF OF THE
;

SAME NATURE ARE UNEQUAL IN DEGREE ( 9C ).

As sleep movements are also manifested by plants growing in tropical climates,

where no injurious nocturnal fall of temperature occurs, the purpose of the sleep
position suggested by DAIIWIN, viz. the protection of the leaves from excessive
cooling, does not hold in all cases. STAHL seeks it also in the lessening of the
formation of dew, and the more energetic transpiration of the unwetted leaves
which would result. Sleep movements are particularly noticeable in Phaseolus,
Trifolium, Jtobinia, Acacia lophantha, Ainicin zygomeris (Fig. 236), Mimosa
pudica, Oxalis, Marsilia, Porlieria while less striking movements take place in
;

many other plants.

Too intense light frequently causes the change from the diurnal position, and
a movement either towards or away from the nocturnal position. The leaflets of
the common Locust (Robinia pscudacacia) are folded downwards at night. In
ordinary diffuse daylight they assume their diurnal, outspread position but, if ;

exposed to the direct rays of the mid-day sun, they turn obliquely upwards.
In many plants ALTERATIONS IN THE INTENSITY OF THE LIGHT ALTER THE
GEOTROPISM OF THE MOTILE ORGANS the sleep movements are then accomplished
;

by the help of geotropic variation movement (Phaseolus, Lupinus) ( ).

97

The change from the diurnal to the nocturnal position continues

for a time to take place, even in constant darkness or prolonged
illumination. The leaves themselves seem to have a tendency to
294 BOTANY

pass at regular intervals from one condition to the other. The

daily periods are the result of the stimulus imparted by the light,
the periodic action of which induces the regular changes of position.

Fie. 236. Amiela zygoma-is, showing <liurn;il anil nocturnal i>osition of leavi--,.

If, however, the external stimulus ceases to operate, the internal dis-
position still continues for some time to give rise to visible after-effects

(p. 261), until finally, from the abnormal conditions, an abnormal

Fio. 23'. Mimosa pvdicv, with leaves in normal, diurnal fosition to the ; rij,'ht, in the position
assumed on stimulation B, inflorescences.
;

state of rigor (light rigor, dark rigor) and symptoms of disease

are manifested ( 98).
Only a few plants respond with pronounced variation movement*
to mechanical irritation (shock, friction, injury). These alone were
SECT, ii PHYSIOLOGY 295

considered irritable plants, so long as only mechanical irritations,

from which visible movements resulted, were regarded as stimuli.

Of irritable plants in this sense, mention has already been made of Dionaea
muscipula (p. 237), leaves, when touched
whose on the upper side, especially if
the bristles are disturbed, fold together. The most familiar example of this
irritability to mechanical stimuli is furnished by Mimosa pudica, a tropical
leguminous shrubby plant, which owes its name
of sensitive plant to its extreme
sensitiveness to contact. The leaves of this plant are doubly compound (Fig. 237).
The four secondary kaf-stalks, to which closely crowded leaflets are attached left
and right, are articulated by well-developed pulvini with the primary leaf-stalks ;

while they, in turn, as well as the leaflets, are similarly provided with motile
organs. Thus all these different parts are capable of independent movement, and
the appearance of the entire leaf becomes, in consequence, greatly modified. In
their unirritated, light position (Fig. 237, on the left) the leaf-stalk is directed
obliquely upwards, while the secondary petioles with their leaflets are extended
almost in one plane. Upon any vibration of the leaf, in favourable conditions of
temperature (25-30 C.) all its parts perform rapid movements.
and moisture,
The and, at the same time, move forward, the secondary
leaflets fold together,

petioles lay themselves laterally together, while the primary leaf-stalk sinks
downwards (Fig. 237, on the right). Leaves thus affected, if left undisturbed,
soon resume their former position.
The behaviour of the leaves is still more remarkable when only a few of the
upon by the stimulus. This is easily demonstrated by holding a
leaflets are acted

burning match near the leaflets of one of the pinnfe. The leaflets directly affected
by the flame fold quickly upwards, and this movement is performed successively
by each pair of leaflets of the pinna until the articulation with the primary leaf-
stalk is reached. The stimulation is then conveyed to the other pinnse, the
which go through the same movement in a reverse order finally, the
leaflets of ;

secondary petioles themselves draw together. Suddenly, when the whole process
seems apparently finished, the main leaf-stalk in turn makes a downward move-
ment. From this leaf the stimulus is able to travel still farther through the
stem, and it may thus induce movement in leaves 50 cm. distant.
The movements of the pulvini are due solely to differences in turgidity which,
as in the case of nyctitropic movements, occur antagonistically in the halves of
the pulvinus. It has been observed that a sudden escape of water into the inter-
cellular spaces takes place out of the cells of the lower or irritable side of the

pulvinus of the primary leaf-stalk the lower surface is that provided with tactile
;

hairs. According to the investigations of HABERLANDT, the conduction of the

stimulus does not appear to be accomplished by the movement of the water thus
discharged, but by the mucilaginous contents of tubular cells which are situated in
the phloem portion of the vascular bundles while FITTING regards the conduction
;

as taking place through living cells. MAC'DouGAL was unable to induce the move-
ments by causing differences in the hydrostatic pressure. Tiie position of an
irritated leaf resembles externally its sleep or nocturnal position, but in reality the

turgor tension of the pulvinus is different (").

Neptunia oleracea, Desmanthus plenus, and Biophytum sensitivum are similar

though less sensitive. Robinia psevdacacia and Omlis acetosella respond to violent
mechanical stimuli by slight but similar movements.

The state of rigor sometimes occurring in motile organs may also

296 BOTANY

be best observed in Mimosa, for, although so sensitive to the action

of external influences, it does not exhibit its irritable movements at
all times. Whenever the temperature of the surrounding air falls
below a certain level, no movements take place, and the whole
plant passes into a condition known as COLD RIGOR, while, on the
other hand, at a temperature of about 40, HEAT RIGOR occurs.
DROUGHT RIGOR is induced, just before withering, by an insufficient
supply of water, and a DARK RIGOR by a prolonged retention in
darkness, probably owing to the derangement of the chloroplasts. In
a vacuum, or on exposure to hydrogen
and other gases chloroform vapour,
coal gas, etc. movement also ceases,

partly on account of insufficient oxygen,

and partly from the actual poisonous
action of the gases themselves. If the
state of rigor is not continued too
long, the original irritability will again
return on the restoration of normal
10
conditions ( ).

The movements of irritability exhibited by

the staminal leaves of some Berberidaceae
(Herberts, Mahonia) and Compositae (Cynareae
and Liguliflorae) bear a certain relation to
those of foliage leaves. The bow-shaped
filaments of the stamens of the Compositae
straighten upon mechanical irritation. As
they frequently contract 10-20 per cent of their
length, the style becomes extended beyond
the anther-tube (Fig. 238). The reduction
FIG. 238. -Asingle flower of Centaurea
{n thej h Qf the filaments isaccompan i e d
jacea with perianth removed. A, sta- . . . . . ,

mens in normal position B, stamens ;

by a moderate increase in their thickness, due
contracted ; c, lower i>art of tubular to the elastic contraction of the cell walls,
perianth ; s, stamens ; a, anther-tube ;
an(j the consequent expulsion of water into
;/, style ; P, pollen. (After PFEFFER, interce ll u l ar spac es. The stamens of
th(j

Berberis and Mahonia are only sensitive to

contact on the inner side near their base, and as their contraction occurs only on
101
the inner side, the anthers are thus brought into contact with the stigma ( ).
of variation movements of carpellary leaves may be seen in the flowers
Examples
of Mimulus, StroUlanthes (Goldfussia), 'Martynia, Torenia, and other plants.
The two lobes of the styles of these flowers fold together when irritated. The
has been
stigmas of Mimulus and Torenia open again shortly unless active pollen
in the flowers of
deposited when they remain in the closed condition. Similarly,
Stylidium, a sudden upward movement of the bent style occurs
when it is
irritated by a touch.
SECT, n PHYSIOLOGY 297

VI. Reproduction

The life is of limited duration.

of every plant Death ensues,
sooner or and the decayed remains form a part of the surface
later,
soil. All existing plants are descended from ancestral forms. A
spontaneous generation of new organisms from lifeless matter does
not, as far as experience teaches, take place, and all existing vegetable
life owes its existence to the capacity inherent in all organisms of

reproducing their kind. Reproduction is accordingly a vital power

which must be exercised by every existing plant species.
It is also evident from the very nature of reproduction that in
the production of new organisms a process of rejuvenation is con-
tinually being carried on. The descendants commence their develop-
ment at a stage long since passed over by the parents.
The formation of independently existing offspring necessitates
also their separation from the parent plant. The formation of a new
bud by a tree is not distinguished as reproduction so long as the bud
remains in connection with the tree as a part of the whole. But if the
bud became separated from the tree and continued its existence as
an independent plant, that would constitute a form of reproduction,
and, in fact, this actually takes place in many plants.
The conditions of the outer world make the still further demand
upon reproduction, that from it a multiplication of the species should
result. As the germs after separation from the mother plant do not
always find the conditions necessary for their development and so, for
the most part, perish, the extinction of the whole species would soon
result if a plant produced but a single germ. That in reproduction
care is taken for the multiplication of the individual in an almost
spendthrift manner, is shown by a consideration of the innumerable
spores produced by a single mushroom, or by a large fern. BP;SSEY
has estimated the number of seeds produced annually by a Poplar
tree at about 28 millions.
REJUVENATION, SEPARATION, and MULTIPLICATION of the individual
are accordingly the essential requisites of reproduction.
These requirements are fulfilled by plants in the most varied
manner. Each great division of the vegetable kingdom has adopted
its own special method ; and each family and genus, or even the
different species, are characterised by some peculiar feature of their
manner of reproduction. Systematic botany is, indeed, essentially
based upon differences in the development of the reproductive organs
and in their functions.
Numerous and varied as the processes of reproduction in the
vegetable kingdom are, they are in reality but modifications of two
different and distinct modes of reproduction.
The simpler of these, or VEGETATIVE REPRODUCTION, consists in
298 BOTANY PART I

the formation of cells or cell-bodies which, after their separation from

the parent plant without undergoing any further change, either
germinate at once, or develop into new organisms after a period of
rest. This mode of reproduction, in which the growth and develop-
ment of the parent plant are directly continued, is also distinguished
as MONOGENETIC, VEGETATIVE, or ASEXUAL reproduction. The
special peculiarities and laws of growth of the parent plant are in
this case directly transmitted to the offspring.
In SEXUAL REPRODUCTION, the second of the two modes of
reproduction, two kinds of reproductive cells each of which carries
the characters of the organism producing it are first formed,
but neither is directly capable of further development, and both
perish in a very short time, unless opportunity is given for their
fusion with each other. Not until one cell (the female) has fully
taken up and become inseparably united with the other cell (the male),
does it acquire the capacity of development and growth. This mode
of reproduction is designated SEXUAL or DIGENETIC reproduction.
The physiological significance of sexual reproduction is not at
once apparent. In many plants the vegetative mode of reproduction
is sufficient to secure the necessary multiplication of the species, so

that plants are able to continue without sexual reproduction. Many

Fungi, for instance, are reproduced only vegetatively ; the cultivated
Banana, many Dioscoreaceae, and varieties of the Grape, Orange,
and the Strawberry, no longer reproduce themselves sexually, but are
propagated solely in a vegetative manner. The Garlic, which forms
small bulbs in place of flowers, the White Lily, and ReunuMuihu
Ficaria, which reproduces itself by root tubers, are hardly able to
produce good seeds, which can, however, be obtained by operative
interference, e.g. on separated inflorescences. They multiply ex-
clusively by asexual methods without suffering any degeneration.
Continued reproduction by vegetative means used to be regarded as
102
necessarily injurious ( ).
Since monogenetic reproduction is sufficient for the preservation
of the species, sexual reproduction must answer some purpose not
attained by the vegetative mode of multiplication, for otherwise it
would be altogether superfluous that the same plant, in addition to
the vegetative, should also possess the sexual form of reproduction,
which is so much more complicated and less certain. Even the
common Moulds, whose vegetative spores (conidia) are very widely
distributed, occasionally develop sexual reproductive cells in specially
formed sexual organs.
In many of the lower plants (Algae and Fungi) it has been shown
that the development of sexual cells is dependent upon definite
external influences. KLEBS has demonstrated, in fact, that it is

possible by regulation of the external conditions (nutrition,

temperature, transpiration, composition of substratum and of the
SECT, u PHYSIOLOGY 299

surrounding medium) to induce them to produce at will either non-

sexual swarm-spores or sexual cells. In many plants unfavourable
external conditions apparently give the impetus to a sexual mode of
reproduction. The sexual product (zygospores of Algae, Phycomy-
cetes) seems better able than the vegetative germs (swarm-spores of
Algae, conidia) to remain a long time at rest, and so withstand the
disastrous effects of an unfavourable environment. No inference can
be drawn, however, from the function of the sexual germs in this
instance concerning 'the necessity for the existence of a sexual, in
addition to a vegetative, mode of reproduction ; for in other cases it
is the vegetative reproductive bodies, as, for example, the spores of
103
Ferns, which are especially equipped for a period of enforced rest ( ).
What makes digenetic reproduction essentially different .from
monogenetic isUNION OF THE SUBSTANCES OF THE PARENTS AND
the
THE CONSEQUENT TRANSMISSION AND BLENDING OF THE PATERNAL
AND MATERNAL PROPERTIES.
It is in this qualitative influence that the chief difference between
sexual and vegetative reproduction is shown. And this may be re-
garded as the special advantage of sexuality. BY VEGETATIVE REPRO-
DUCTION^ THE QUANTITATIVE MULTIPLICATION OF THE INDIVIDUAL IS
SECURED, WHILE BY SEXUAL REPRODUCTION A QUALITATIVE INFLUENCE
IS EXERTED. THE VEGETATIVELY PRODUCED PROGENY CONSIST OF
UNMIXED DESCENDANTS THE SEXUALLY PRODUCED OFFSPRING, ON
;
"

THE OTHER HAND, ARE THE RESULT OF A BLENDING OF THE PARENTS.

In vegetative multiplication the complex of properties unfolded
in the descendants does not as a rule differ from that possessed by
the parent form. By vegetative multiplication all the varieties and
races of cultivated plants, the characters of which do not come true
by seed, are maintained. Variation may, however, occur among the
vegetatively produced progeny. The occurrence of this may be
compared to bud- variation, in which a single bud on a tree may give
rise to a branch deviating in colour arid form from the type ; such a
bud-variation may, years after, return suddenly to the characters
of the typical form. In the same way variations arise among
vegetatively produced plants, a remarkable example of which, .accord-
ing to BEYERINCK, is afforded by the Bacteria, the reproduction of
which takes place exclusively by division into two ( 104 ).
The sexually produced offspring, on the other hand, endowed
with the properties of the father, can never be identical with the
mother-plant, but possess the properties of both parents. When these
are divergent they frequently play very different parts in the
descendants, some (dominant) characters appearing conspicuously,
while others (recessive characters) become less marked or remain
completely latent. la this way the descendants do not exhibit a
uniform mean between the parents, but some may resemble the father,
others the mother. These relations determine the character of the
300 BOTANY PART T

sexually produced descendants (cf. p. 313

Variations appearing
ff.).
an absolutely dominating
in single individuals will, unless they are of
character, become modified and ultimately lost by crossing with
ordinary individuals. In such a case sexual reproduction tends to
maintain the constancy of the species. In other cases, as when one
parent possesses new and dominant characters or when both parents
tend to vary in the same direction, the deviation from the ancestral
form may be maintained or increased by sexual reproduction.
The great tendency to variation
commonly exhibited by hybrids
(p. 313) illustrates howthe equilibrium of the complex of properties
of a sexually produced individual is affected by divergent parental
tendencies. But, even as a result of ordinary fertilisation, not only
small and readily disappearing variations (fluctuating variations) but
sometimes more striking ones occur, in which the offspring differs so
strongly from the parents in characters, which can be inherited, that it
appears to be a new species or sub-species. Of such petites esptces of
Draba verna some two hundred are known. In such sudden variations
(the occurrence of which v. KOLLIKER, and with him KORSCHINSKV,
term heterogenesis, while DE VRIES more recently calls it mutation)
these authors seek the starting-points of the origin of new species.
This would occur when a particular species passes, from unknown
causes, into a period of mutation such as DE VRIES demonstrated
experimentally in Oenoihera Lamarckiana. KORSCHINSKY collected a
number examples of heterogenesis, of which Chelidonium
of historical
laciniatum, which appeared in a garden at Heidelberg in 1590, and
Capsella Heegeri, SOLMS, which only recently appeared, will serve as
105
examples ( ).

The fluctuating variations which largely determine the valuable characters of

economic plants (e.g. the high percentage of sugar in the Sugar Beet) are in con-
trast to the mutations not fixed on inheritance. Careful and continued selection of
the varying progeny is thus necessary to maintain the required standard of the race.

The experience of cultivators and the recent experimental work

on lower plants carried out by KLEBS show that different groupings
of the internal and external conditions of life favour reproduction
and ordinary growth respectively. In fact growth and reproduction
frequently though not always appear to be mutually exclusive.

Vegetative Reproduction

Vegetative reproduction, the .purely quantitative character of

which as a mere process of multiplication has been emphasised, exists
generally throughout the vegetable kingdom, and but few plants, e.g.
some of the Conifers and Palms, are altogether devoid of it. Mention
has already been made in considering artificial propagation that,
from the separate parts or single cells, or even from the naked
protoplasts (Siphoneae) of many plants, the regeneration of a new and
 PHYSIOLOGY 301

perfect individual may ensue. In vegetative reproduction the process

is similar except that the separation of the part from tBe parent plant

is an organic one, occurring in the natural course of development.

The vegetative form of reproduction is manifested in various aspects,

and may be distinguished as a multiplication by means of multicellular
vegetative bodies (buds, gemmae), or by single cells (spores).
Multiplication by Multieellular Vegetative Bodies (Budding )
1

often consists merely in the separation of lateral shoots, or in a

division of a single plant into several. In this way the lateral
shoots of Azolla, through the death and disruption of the older
parts of the parent axis, become separated from one another and
continue their growth as independent plants ; similarly, separate
plants originate from the vegetative body of the Duckweed (Lemna).
Multiplication by stolons, rhizomes, and tubers results in a similar
formation of independently existing plants. As may be seen in the
Strawberry, the Bugle (Ajuga, reptans) and numerous other plants,
the shoots produced from many of the axillary buds of the widely
outstretched stolons take root and form new plants. In cases where
the runners themselves eventually die, the parent plant becomes finally
surrounded by a colony of entirely independent plants. Instead
of forming runners, the single tuber may divide (Corydalis solida),
and in this way give rise to two, four, or more new tubers. New
bulbs are produced in the leaf-axils of the bud-scales of bulbs, while
brood buds (bulbils, gemmae) are frequently developed on aerial
vegetative organs.

Bulbils are found on the inflorescence in the place of the flowers in many species
of Allium, in the grass Poa bulbifera, and also in Polygonum viviparum. In
Lilium bulbiferum, Dentaria bulbifera, etc., the bulbs in the axils of the leaves are
specially constructed with a view to detachment from the parent plant (Fig. 22).
The swollen leaves contain reserve food material, and frequently develop roots before
falling from the plant. In Ranunculus Ficaria the roots of the axillary buds are
full of reserve food material,and resemble grains of corn. When the plant dies the
bulbils remain on the ground, and have given rise to the fable of showers of grain.
Bulbils or gemmae are met with also among the Mosses, Liverworts, and Ferns. The
winter buds of many water plants (Hydrocharis, Utricularia, Lemna, etc.) have a
peculiar biological significance. They are formed in the autumn, and sink to the
bottom of the water ;
in the succeeding spring they rise to the surface and form
new plants.
By vegetative multiplication higher plants can annually give rise to individuals
which are strong and capable of flowering and fruiting. The seedlings of sucli
plants, on the other hand, often require to grow for several years before the
capacity of sexual reproduction is attained (bulbous plants, Hop, etc.).
In addition to the instances just cited, in which the vegetative reproductive
bodies take their origin from points where lateral shoots are normally formed, they
may also appear in places where no shoots are normally developed. Thus the
adventitious formations often found on leaves, particularly on the leaf-blades,
serve the purpose of reproduction. Just as the leaves of Begonia, Drosero, etc..
302 BOTANY PART I

after they have been cut off, are able to give rise to new plants, in other cases the
leaves jxmess this power while still growing on the parent plant. Some ferns
afford specially characteristic examples of this (Asplenium decussatum, A.
Fabianum, A. bulbiferum, A. viviparum) ; adventitious buds are produced on their
laminae, developing into small rooted plants, which then fall off and complete their
development (Fig. 239). The adventitious buds of Cystopteris bulbifera take the
form of bulbils with small swollen leaves. Adventitious plantlets are frequently
formed also on the leaves of Cardamine pratcnsis, and Cardamine amara manifests
a similar tendency. One of the best-known examples of such adventitious forma-
tions is afforded by the leaves of the tropical Bryophyllum, in the marginal indi-ntu-
tions of which the brood plantlets develop in great numbers. Even the force of the
wind is sufficient to detach the leaflets of Bryophyllum from the plant. Gemmae
are abundantly produced on the thallus of many Hepaticae (Marchantia, Lunularia),
and by their continuous growth the gemma cups (Figs. 381, 382) are always kt-pt
well filled.

One of the most interesting cases of adventitious budding is the formation of

FIG. 230. AfjilenluniFabianum. A young plant (T), with leaves and roots (W),
has sprung from the leaf (J/) of tin- nldi'r plant.

adventitious embryos in ovules ; in most instances this leads to POLYEMBRYONY,

i.e. the presence of several embryos within the one seed. In addition to cases in
which the additional embryos cannot be strictly regarded as adventitious but
arise by fusion of ovules, by division of the nucellus within the integments, or by
the existence of a number of embryo-sacs in one nucellus, normal ovules with only
one embryo-sac may produce a number of embryos. In the latter case the embryo-
sac may in rare instances (Santalum, Sinninyia) contain two egg-cells, or a

dichotomy of the embryo or its suspensor may occur. As a rule the condition is
due to adjoining vegetative cells growing into the embryo-sac and there develop-
ing like sexually produced embryos. These adventitious embryos, which were
first discovered by Strasburger, usually originate from cells of the nucellus (Funkia

ovata (Fig. 240), Euonymus latifolia, Citrus aurantium, Nothoscordon fragrans,

Mangifera indica, Coelebogyne illicifolia, Clusia alba, Opuntia vulgarc, Euphorbia
dulcis, Colchicum autumnale) more rarely from the inner integuments (Allln,i>
odorum). In many other plants (Gnetum, Ulmus) embryos may develop from the
synergidse or the antipodal cells. At the same time the egg-cell previously
existing in the embryo-sac is able to continue its development after fertilisation,
but usually prevented from so doing by the adventitious embryos.
is The seeds
in such cases would no longer contain the products of sexual reproduction, but
SECT. II PHYSIOLOGY 303

would be degraded to organs of vegetative multiplication. The adventitious

germs in the polyembryonic seed are, however, so far dependent upon sexual
reproduction, that for the most part they only attain their development in case
pollination has previously taken place but in Coelebogyne, one of the Australian
;

Euphorbiaceae, of which usually only female specimens are found in cultivation,

and in Bcdanophora elowjata and Elatostema acuminatum according to TREUB,
and Bal. ylobosa according to LOTSY, the adventitious germs develop without the
stimulus of fertilisation. These plants accordingly afford examples of APOGAMY,
or of the substitution of a vegetative for a sexual mode of reproduction, such as
/M-i-urs in certain ferns.

In the apogamous ferns vegetatively produced plants arise on the prothallus

in the position of the sexual organs. This is found in different degrees in
Athyriurnjilixfcmina, var. cristata, As-
pidium falcatum, Todea africana, Pteris
cretica, and in Ncphrodiwn pseudo-mas.
var. polydactyla. In the latter examples
the sexual organs are no longer formed,
although the young plants arise, by a
vegetative process of budding, from
exactly the same part of the prothallium
where the archegonia would have been
developed. FARMER, MOORE, and DIG BY
have shown that in Nephrodiumfae origin
of the apogamous bud is preceded by fusion
of nuclei of vegetative cells of the pro-
thallus. In the case of Aspidium filix
mas. var. cristatum, etc., the apogamy
seems to have resulted from cultivation.
In a broad sense the development of
bulbils in the place of flowers, in the

species of Allium, might be considered Kiu. Vegetative formation of embryos in

:i4u.

as an example of apogamy ( 106 ). Funkia ovata (Hosta coerulea) by the budding

The cases of of the nucleus ; n, nucellus with cells in
parthenogenesis de-
scribed in recent years process of forming the rudiments (ae) of the
among the higher adventitious embryos S, synergidae ; E, egg.
;

plants (Compositae, Alchemilla, Thalic- cell, in the lower figure developing into a sexu-
Irum, firyonia ?) may be regarded as ally-produced embryo ; if, inner integument
examples of a further peculiar type of
apogamy. The ovum develops into an embryo without fertilisation, but since
the reduction division has been omitted from the processes leading to its differ-
entiation the egg cell has in these cases lost the characteristics of a female
sexual cell and corresponds to a purely vegetative cell. The same will probably
be found to be the case in Wikstroemia indica, Ficus hirta, and Chara crinita which
are also parthenogenetic 107
( ).

In some cryptogamic plants (Marsilia,

Saprolegnia) as among
lower animals, true parthenogenesis occurs. LOEB found that solu-
tions which withdrew water
(MgCL,, other salts, sugar, urea) could
stimulate the ovum to parthenogenetic development. WINKLER used
extractives derived from the sperm, while NATHANSOHN showed that
sometimes a rise of temperature could interrupt the
resting state of
an ovum awaiting fertilisation and lead to a
parthenogenetic develop-
304 BOTANY PAKT I

ment. JICKELI regards the common feature of the various causes

which lead to parthenogenesis to be that they are injurious influences,
108
the effect of which, however, must not be carried too far ( ).
Vegetative Multiplication by Single Cells (Spores). As iu the case of multi-
cellular vegetative bodies, multiplication can be effected also through the .separa-
tion of single cells. Strictly .speaking, this manner of multiplication actually
takes whenever a division of the vegetative body occurs in unicellular
place
Bacteria, Fungi, and Algae. which serve the purpose of vegetative reproduc*
Cells

tion, and have a special form and method of development (spores, fouidia, swarm-
spores,gemmae), are first met with in the higher Cryptogams. They are frequently
formed in special organs or receptacles (sporangia, conidiophores, fruit bodies).
As a Fern-plant occasionally arises directly from the tissue of the prothallu.s
without the intervention of the sexual act, so also spore formation is occasionally
omitted, and the prothallus arises vegetatively from the leaf of the sporophyte
(apospory in varieties of Athyrium, Jspuliitin, Aspleniuindimorphum, Nephrodium
108a
2)seudo-mas. var. cristata apospora) ( ).

Sexual Reproduction
For the purpose of sexual reproduction two kinds of cells, male
and female, are produced. Although neither alone is as a rule
capable of development, the actual reproductive body is formed by
the fusion into one cell of two such sexually differentiated cells.
The elements of the male and female nuclei remain for a longer or
shorter time distinguishable in the nucleus resulting from their
10J
fusion ( >).

In fertilisation, as a rule, two uninucleate cells fuse, even when the vegetative
protoplasts are multinucleate.
In Albugo Bliti, A. portulacae and A. tragopogcniis,
however, STEVENS found that the numerous nuclei of the ovum fused with an equal
number of sperm nuclei. Probably other cases of this kind may be found.
The further development of non -nucleated fragments of the egg when united
with a spermatozoid has been shown to occur among animals by O. and R. HERTWIO,
BOVEKI, and DELA.GE. This phenomenon which is termed MEROGONY has also been
observed by WIXKLEII iu Cystoscira barbatn, one of the Fucaceae ( no ).
In connection with parthenogenesis it was noted above that the incapability
of further development which characterises the unfertilised ovum may be over-
come by other means than copulation with the male cell KLEBS had already ;

shown this to be the case with the gametes of certain Algae. It is thus necessary
to distinguish in fertilisation between a stimulus, which removes the arrest laid
on the further development of the ovum, and the cell fusion, which influences
the natui'e of the resulting organism (amphimixis). The two influences are united
in the case of natural fertilisation (
m ).

As thus necessary in sexual reproduction not only to provide

it is

for the production of male and female cells, but also to ensure their
union, it becomes at once evident that, for sexual reproduction, the
organs must have a different structure than if they were designed
solely for vegetative activity. The sexual organs accordingly often
exhibit a special and peculiar form, and differ materially in appear-
ance from the vegetative parts of a plant.
 PHYSIOLOGY 305

The Union of Sexual Cells (Fertilisation). Leaving out of con-

sideration the necessary contributory external contrivances, fertilisa-
tion is accomplished by means of a chemotactic or chemotropic stimulus

(pp. 266, 286). It is generally the non-motile egg-cells or the female

sexual organ which exert an attractive influence upon the motile male
cells. When, however, there is no difference in the external fqrm of
the male and female cells, then both are usually motile, and the attrac-
tion seems to be exerted mutually. This is probably the case with
the motile and externally similar sexual cells (GAMETES) of the lower
Cryptogams, particularly of the Algae (Fig. 97). In the conjugation
of the Corijugatae, however, although both sexual cells are externally
alike, one cell alone is usually motile, and passes through the connect-
ing canal to the other. The capacity of the male cells for independent
movement is common to most Algae, with the exception of the
Florideae, in which the non-ciliated male cells are passively conveyed
to the female organ by the water. Throughout the whole group of
the higher Cryptogams, and in a few Gymnosperms, the male cells are
motile spermatozoids, capable of seeking out the non-motile egg-cells
concealed within the archegonia. But in the sexually differentiated
Fungi the male substance usually remains enclosed in special hyphae
which press themselves close against the female organs, and, by the
perforation of the intervening cell wall, the fusion of their contents
is rendered
possible. The fertilisation of the Phanerogams is accom-
panied by a perforation of the intervening cell walls similar to that
which occurs in the Fungi. In this case the male cell is enclosed
within the microspore (pollen grain) ; the female, as a naked egg-cell,
is included in the
megaspore (embryo-sac), which in turn lies in the
ovule, and in the Angiosperms the ovule is again enclosed within the
ovary. The double-walled pollen grains possess no independent power
of movement, but are conveyed to the female sexual organs by the
assistance of external agencies (animals, currents of air or water).
The pollen grain then grows out into a tube which is acted upon by
chemotropic (including hydrotropic and aerotropic) influences, and
grows like a fungal filament through the tissues of the ovary and
ovule until it penetrates to the egg-cell in the embryo-sac the ;

union of the sexual cells is then easily effected (Fig. 99).

Recent researches have shown that in addition to the iusiou of the generative
cell with the ovum the second generative cell frequently fuses with the nucleus of
the embryo-sac. This has been termed DOUBLE-FEKTII.ISATION. The endosperm,
which arises from the resulting nucleus, is thus, like the embryo itself, a product of
fusion. This explains the "hybrid character of the endosperm in the phenomenon of
XI.XIA. In tin-so cases characters of the pollen-parent appear in the ripening seed
and not, as isusual, only in the descendants of the union. This is especially well
shown in the maize when, e.g. blue- and yellow-fruited races or a race the fruits of
which contain sugar and one in which they contain starch are interbred. It is
uncertain what significance is to be attached to this so-called double fertilisation ;

X
306 BOTANY P\KT i

possibly the explanation is that the embryo-sac nucleus, as a sister nucleus to that
of the ovum, i-\nvi~r> a similar chemotactic influence, and thus attracts the male
generative cell, the fusion 1 icing of secondary importance since the endosperm is

destined to be used as food by the embryo C - 1

.

While one healthy pollen grain should be able to fertilise one

healthy ovule, experience shows that better results follow from inure
liberal pollination of the stigma. This depends, according to
COKUENS, on the fact that not every pollen grain (and not every
ovule) is good. Thus in Mvrabttis jalapa for each fertile pollen grain
four are found infertile, and for every three good ovules there is one
bad one. For Miralills loxgijlora the corresponding ratios are 1 3 :

3
and 1 : 1
( ).

To render certain the accomplishment of this POLLINATION, or

conveyance of the pollen to the female sexual organs, special and
often complicated contrivances are made use of by the different
Phanerogams, according to the means of conveyance upon which they
are dependent.
Plants, the pollen of which is carried
by wind, are designated
.\\KMOPHILOUS. As this method of conveyance depends upon the
chance of wind direction, the production of an enormous amount of
pollen characterises wind-fertilised plants. According to Holden a
medium-sized plant of maize produces about 50,000,000 pollen grains.

Such enormous quantities of pollen are often taken up from pine forests by the
wind that clouds of pollen fill the air. The surface of Lake Constance in spring is
so thickly covered with pollen that it is coloured yellow (" the lake blooms," it is

then and in the Norwegian fiords, at a depth of 200 fathoms, the pollen of
said),
Conifers, according to F. C. NOLL, forms for a time the principal nourishment of
a Rhizopod (Saccamina).
The male flowers of such anemophilous plants are accordingly either freely
exposed to the wind in Catkins (Couiferae, Querciflorae), or the versatile anthers,
as in the Grasses, depend from long, lightly-swaying filaments. The pollen grains
themselves do not stick together but escape from the opened anthers in the form of
fine powder. The pollen grains of many Conifers are rendered extremely buoyant
and easy of conveyance by the wind by two sac-like protrusions of the exinc. In
some anemophilous plants the pollen is discharged by the sudden extension of the
filaments, previously rolled up in the bud (Urticaceae, e.g. Pilca), or by the hygro-
scopic tension of the anthers. The female organs are also often specially adapted
for the attachment of the pollen thus floating in the air. The stigmas either spread
out like a brush (Corylus), or are finely feathered or provided with hairs (Grasses,
Walnut), or drawn out into long threads (Indian Corn). In the Conifers, with
freely exposed ovules, the grains of pollen are caught and retained, in a drop of
fluid exuded from the micropyle, into which they are gradually drawn as the fluid
dries up. In other Conifers whose ovules are concealed in the cone of the female
inflorescence, processes of the integument catch the pollen and conduct it to the
sticky opening of the young ovules.

For the fertilisation of the higher plants, the presence of water is

not so essential as it is for most Cryptogams. Only a few submerged
SECT, ii PHYSIOLOGY 307

Phanerogams make use of the agency of water for effecting their

pollination, and are, on that account, termed HYDROPHILOUS PLANTS.
The pollen of the submerged Zostera exhibits certain peculiarities, distinctly
referable to the necessity of effecting fertilisation under water. It does not form
round grains, but in their place elongated thread-like filaments devoid of an exine,
which, as they have the same specific weight as the surrounding water, are easily
set in motion by the slightest currents, and are thus brought into contact with the

stigmas. In the case of the submerged water plants, Vallisneria, Elodea, and
species of Enhalus, found in the Indian Ocean, the pollination is accomplished on
the surface of the water. Thus, for example, the male flowers of Vallisneria, after
separating from the parent plant, rise to the surface of the water, where they open
and float like little boats to the female flowers, which, by the elongation of their
spirally coiled flower-stalks, ascend, at the same time, to the surface of the water,
114
only to become again submerged after fertilisation ( ).

In the great majority of Phanerogams pollination is effected by

means of animals. By enticing in various ways insects, birds, bats,
or snails, plants are enabled not only to utilise the transporting
power but also the intelligence of animals in 'the service of pollen-
conveyance. The pollination is then no longer left to chance ; and
as the transport of pollen to the sexual organs becomes more assured,
the necessity for its formation in such enormous quantities as in
anemophilous plants is obviated. For the most part, such plants
are adapted to POLLINATION BY INSECTS (ENTOMOPHILY). For
their nourishment, plants offer not only the sugary sap, which, as
nectar, is excreted from different parts of the flowers, but also the
pollen itself, which furnishes a nitrogenous food material and,
together with the honey, is kneaded by bees into bee-bread. As
additional means of enticement, and to attract animals from a
distance to the nectar offered by the sexual organs, special perfumes
and conspicuous colours have also been developed. The ATTRACTIVE
APPARATUS of plants is generally formed by the coloured floral
leaves ; by the outer floral leaves or calyx (Nigella, Aconitum), or by
the perianth (Lily, Tulip), or by flowers specially modified for this
purpose (outer florets of the Compositae, Umbelliferae, and Viburnum
opulus ; scented flowers of Eenanthera Lowii). Bracts and portions of
the axis which do not belong strictly to the flower occasionally serve
the same function as in the circumfloral attractive apparatus of the
Euphorbiaceae or the extra-floral attractive apparatus of Astrantia
major, Salvia, Melampyrum, Dalechampia, Bougainvillea spectabilis, and
the Aroideae.
The pollen entomophilous, in contrast to that of the
of the
anemophilous plants, not a dry powder, but its grains are stuck
is

together with an oily or mucilaginous fluid ; in other cases, they are

held together by their rough outer surfaces or by means of viscid
threads (Oenothera\ and can only be removed from the anthers by
animals. The structure of the flower is so contrived, as CHRISTIAN
X l
308 BOTANY PART i

CONRAD SPRENGEL first pointed out in 1793 in his famous work on

the structure and fertilisation of flowers ("Das entdeckte Geheimniss

der Natur im Bau und in der Befruchtung der Blumen "), that the
pollen grains must necessarily become attached to certain parts of
the body of the animal visiting it in search of food, and so be con-
veyed to the sticky or hairy stigma of other flowers. The variety of
means employed to secure pollination, and the wonderful adaptation
shown by the flowers to the form and habits of different insects, are
most remarkable.

In addition to the stimulus of hunger, plants utilise the reproductive instinct

of insects for securing their pollination. Not a few plants (Stapelia, Anstolochia,
and members of the Araceae), by the unnatural colour of their flowers, combined
with a strong carrion-like stench, induce carrion-flies to visit them and deposit
their eggs in so doing they effect, at the same time, the pollination of the flowers.
;

In the well-known hollow, pear-shaped inflorescences of the Fig (Ficus carica,

Fig. 575), there occur, in addition to long-styled female flowers that produce seeds,
similar gall-flowers with short styles. In each of the latter a single egg is laid by
the Gall -wasp (Blastophaga), which in effecting this pollinates the fertile flowers
with pollen carried froin the male inflorescence (the Caprificus). The large white
flowers of Yucca (Fig. 493) are exclusively pollinated by the Yucca moth (Pronuba).
The moth escapes from the pupa in the soil at the time of flowering of Yucca and
introduces its eggs into the ovary by means of the style ;
in doing this it carries

pollen to the stigma. The larvae of the moth consume a proportion of the ovules
in the ovary, but without the agency of the moth no seeds would be developed, as
the sterility of the plant in cultivation shows.
In South America the humni ing-birds are especially active in the conveyance
of pollen, as they seek for insects in the flowers ; a starling visits the flowers of
species of Puya to drink the watery nectar. In the Old World the honey-birds
play a similar part. MARLOTH enumerates about forty ornithophilous plants from
the South African flora these are mostly pollinated by species of Nectarinia.
;

Species of Feijoa have sweet succulent perianth leaves to attract the birds, which
serve to convey the pollen.
Besides these ORNITHOPHILOUS plants there are a few visited by Bats
(CHIROPTEROPHILOUS) thus the dioecious Pandanaceous plant Freycinetia is
;

pollinated by a Flying Fox (Pteropus), which eats the inner bracts.

Pollination in some cases is effected by means of snails (MALACOPHILOUS
PLANTS). To their instrumentality the flowers of Calla palustris, Chryso-
splenium, and also the half-buried flowers of the well-known Aspidistra owe their
pollination (
m ).

Self- and Cross-Fertilisation. It has already been pointed out

that it is in contrast to the vegetative mode
by sexual reproduction,
of multiplication, that qualitative modifications are effected. Such
qualitative changes are best attained when the sexual cells are derived
from different individuals. It is in accordance with this same principle
that, in the sexual reproduction of plants, varied and often complicated
contrivances are manifested, which conduce to CROSS-FERTILISATION
{union between sexual cells of different individuals), even when the
 PHYSIOLOGY 309

individuals themselves are HERMAPHRODITE and possess two kinds of

sexual organs, as in the case of the majority of Phanerogams.
As, however, self-fertilisation takes place also in a number of
plants, either regularly or as a makeshift, it is evident that Avhat-
ever may be the advantage derived from a union of two distinct
individuals, it is no more essential for sexual reproduction than
for vegetative multiplication. Self-pollination, although regularly
occurring, frequently fails to occasion self-fertilisation, as often the
pollen will not develop pollen-tubes on the stigmas of the flower
(self-sterile) by which it was produced, but only on those of different
flowers (Secale cereale, Corydalis cava and some Crucif erae, Lobelia fulgens,
116
Verbascum nigrum, etc.) ( ).

The antipathy between the sexual organs of the same flower, in certain plants,
so greatly exceeds thebounds of indifference that they act upon each other as

poisons. Thus, for example, it is known of certain Orchids that pollination with
their own pollen causes the death of the flower, while in other cases the pollen is
killed in a short time by the stigmatic fluid of the same flower.

In other instances, self-fertilisation occurs where cross-pollination

either is not effected, or else as an alternative to it (Wheat, Barley,

Canna, Viola species, Linum usitatissimum, etc.). By many plants, in

"
addition to the large " chasmogamous flowers adapted to insect
pollination, small, inconspicuous flowers are produced which, usually
concealed underg'round or by the lower leaves, never open, and only
bear seeds which have been produced by self-fertilisation. In such
flowers the stamens no longer open, the pollen-tubes growing through
the wall of the anther to reach the stigma. In some plants the
majority of the seeds are derived from such CLEISTOGAMOUS flowers
(Viola), and sometimes their seeds alone are fruitful (Polycarpum
tetraphyllum possesses only cleistogamous flowers). As the greater
number of such plants, however, in addition to the seeds of the self-
fertilised small cleistogamous flowers, produce seeds resulting from
the cross-fertilisation effected in the larger flowers (Impatiens noli-
tangere, species of Lamium, Specularia perfoliata, Stellaria, Juncus
bufonius, etc.), the ancestral plants of the cleistogamous generations,
as well as their descendants, have, at least, the opportunity for
cross-fertilisation open to them.

Special contrivances for ensuring the crossing of the sexual cells,

particularly by preventing self-pollination, are found to exist through-
out the whole vegetable kingdom.
Self-pollination is effectually avoided when the plants are
most
unisexual, that is when
there are both male and female plants.
Such DIOECIOUS plants exist in almost all classes of plants from the
lower Cryptogams to the most highly developed Phanerogams (certain
Mucorineae, many of the lower Algae, species of Fucus, Marchantia,
Polytrichum, Equisetaceae, Taxus, Hemp, Hops, Date-Palm, etc.) In
X 2
310 BOTANY

MONCECIOUS plants (other Mucorineae, Algae, species of Carer, liicinus,

Box, Coniferae, Cupuliferae, etc.) the male and female organs occur
on different flowers, but the flowers are borne on the same plants.
The fertilisation between different flowers is thus secured but even ;

here crossing with other individuals is, for the most part, assured by
dichogamy.
The term DICHOGAMY is used to denote the fact that the male and female sexual
organs attain their maturity at different times. When either the male or female
sexual organ matures before the other, the self-pollination of morphologically
hermaphrodite flowers is avoided and crossing ensured. Both hermaphrodism and
moncecism are more advantageous than dicecism, as
all the individual plants in such cases are able to
produce seeds while in dioecious plants the male
;

flowers cannot be utilised for the direct production of

seeds. Dichogamy secures crossing in such a simple
Tiianner. and is so easily attained by hermaphrodite
plants, that it of very general occurrence in the
is

vegetable kingdom. According to the priority of the

maturity of their sexual organs, plants are designated
PROTANDHOUS or PROTOGYNOU8.
PROTANDRY, the earlier maturing of the male
sexual organs, is the more frequent form of dichogamy.
It occurs in the flowers of the Geraniaceae, Campanu-

laceae, Compositae, Lobeliaceae, Umbelliferae, Mal-

vaceae, etc. The anthers, in this case, open and dis-
charge their pollen at a time when the stigmas
of the same flowers are still imperfectly developed
and not ready for pollination. Accordingly, PKO-
TANDROUS FLOWEKS CAN ONLY BE FERTILISED BY THE
POLLEN OF YOUNGER FLOWERS.
FIG. 241. Inflorescence of Plan- In the less frequent PROTOGYNY the female sexual
tago media with protogynous organs are ready for fertilisation before the pollen
flowers. The upper, still closed
o f the same flowers is ripe, and the' stigma is usually
flowers (?) have protruding
r uinated and
wit hered before the pollen is shed so ;
styles; the lower (<?) have
lost th.-ir styles, and disclose tnat thePROTOGYNOUS FLOWERS MUST BE FERTILISED
their elongated stain. MIS. BY THE POLLEN OF OLDER FLOWERS (Anthoxanthum
odoratum, Luzula pilosa, Scrophularia nodosa, Aris-
tolochia clematitis, Helleborus, Magnolia, Plantago, Fig. 241).
A still morecomplicated
because in-
method of effecting cross -fertilisation,
differences of structure, results from
volving also morphological and anatomical
HETEROSTYLY, or the peculiarity of some species of plants of producing stigmas
and anthers which vary in height in different individuals of the same species.
In contrast to the unlimited possibility of crossing in other hermaphrodite
flowers (panmixia), there is here a limitation to certain classes of individuals of
the species. A good example of heterostyled flowers is afforded by the Chinese
Primrose (Fig. 242). This plant has two forms of flowers, long-styled (L) and
short-styled (K), while the positions of the stigmas
and anthers in the two kinds
of flowers are exactly reversed. The pollen grains of the short-styled flowers,
moreover, are larger, and the stigmatic papillae shorter, than in those with the
longer styles (p, P, and n, N). The purpose of such morphological and anatomical
SECT, ii PHYSIOLOGY 311

differences existingbetween flowers of the same species was first understood after
they were discovered by DARWIN to be a contrivance for cross-pollination. Ferti-
lisation is most successful in such cases when the pollination of the stigmas is
" "
effected by the pollen of anthers correspondingly situated. By such a legitimate
fertilisation, more and better seeds are produced than by "illegitimate" fertilisa-
tion, and in some cases (Linum perenne, Fagopyrum csculentum) legitimate
fertilisation alone is productive. Legitimate fertilisation is rendered more certain
by the fact that insects in visiting the flowers touch correspondingly placed sexual
organs with the same portions of their body. The flowers of Primroses have styles
of two different lengths (DIMORPHIC HETEROSTYLY) the same peculiarity is
;

exhibited by Piilmonaria, Hottonia, Fagopyrum, Linum. There are also flowers

with TRIMORPHIC HETEROSTYLY (Lythrum Salicaria, and some species of Oxalis),
in which there are two circles of stamens and three variations in the height of the
stigmas and anthers.

Fio. 242. Primula sineiisis: two heterostyled flowersj'from- different plants. L, Long-styled;
K, short-styled flowers G, style S, anthers
; ; P, pollen-grains, and N, stigmatic papilke of
;i

the 'long-styled form; p and n, pollen-grains and- stij-'inatic papillae of the short-styled form.
(P, N, p, n, x lio.)

The different forms of heterostylic plants do not only differ in the length of the
style and stamens, and in the colour and of the pollen grains, but according
size,
to ERRERA commonly exhibit differences in form of leaf, in the size and colouring of
the flowers, and in the weight of the seeds.
is made mechanically impossible,
In a great number of flowers self-pollination
as their own
pollen is prevented by the respective positions of the sexual organs
from coming in contact with the stigma (HERCOGAMY). In the Iris, for example,
the anthers are sheltered under the branched petaloid style, upon whose lip-like
stigma no pollen can come, unless through the agency of insects, and in the
protogynous flower of Aristolochin dcrnatitis pollination from the anthers,
which occupy a lower position on the column, is prevented. The conveyance of
pollen from the older to the younger flowers is effected in Aristolochia by small
insects. The flowers at first stand upright with a widely .opened mouth (Fig. 243 /)
and in this condition the insects can easily push past the downwardly directed
hairs clothing the tubular portion of the corolla and reachjthe dilated portion below.
Their exit is, however, pi-evented by the hairs until the stigma has withered and
the anthers have shed their pollen. When this has taken place (Fig. 243 IT) the
hairs dry up and the insects covered with pollen can make their way out and
312 BOTANY

convey the pollen to the receptive stigmas of younger flowers. In the Orchidaceae
and Asclepiadaceae self-pollination is rendered impossible both by the nature of
tlie pollen masses and
by their position.
A complicated form of structural
contrivance, by means of which cross-

pollination is secured, may be seen in

a flower of Salvia pratensis (Fig. 244).
The anthers of this flower are concealed
in the upper lip of the corolla, from
which the style, with its bilobed
stigma, projects. When a bumble-bee
visits the flower in search of honey,
it must first with its
proboscis push
out of the way the small plate (s),
formed of two sterile anther halves
grown together. These are situated
at the ends of the short arms of the
connectives (c), which are so elongated
that they might easily be mistaken
for the filaments (/) of the stamens.
The fertile -anther halves are situated
at the other ends of the connectives,
and are thus brought in contact with
Fio. 243. -Flowers of Aristolochia dematitis cut the hairy back of the bumble-bee when
through longitudinally. /, Young flower in which it pushes against the plate at the short
the stigma (N) is receptive and the stamens (S) e nds of the lever-like connectives. The
have not yet opened II, Older flower with the
;

r Ren t, mg attached to the bee will

stamens opened, the stigma withered, and the , ,
. . . . , . ,

hairs on the coroila dried up. (x _>.)

back by the forked
be brushed oil its

stigma borne on the elongated style

of an older flower (Fig. 244 //). Good examples of hercogamous flowers are afforded
by the Papilonaceae, by Kalmia, whose anthers are held in pockets of the corolla
and are only released when touched by a visiting insect (Fig. 245), by Vincu, etc.

Fio. 244. Pollination of Salvia pratensis. 1, flower visited l.y a l,unil>le-l>ee, showing tho projec-
tion of the curved connective from the helmet-shaped upper lip, and the de]>ositi<m <>t !]
pollen on the back of the bumble-bee; 2, older flower, with connective
drawn Uirk, and
the upper
elongated style ; 4, the staminal apparatus at rest, with connective enclosed within
lip ; 3, the same, when disturbed by the entrance of the proboscis of the
bee in the direction of
the arrow /, filament ; e, connective s, the obstructing halt of the anther.
; ;

Some that are purely

plants bear in addition to hermaphrodite flowers others
female. are termed and the distinction is made between
They polygamous,
 PHYSIOLOGY 313

gynodujecious and gynomonoecious forms. In the latter the female flowers are
borne on the same individual as the hermaphrodite flowers (Aesculus, etc.) ; in
the former they are on distinct individuals
(Silene inflata, Erodium cicutarium, Satureja).

Hybridisation. The union of two

sexual cells is, as a rule, only possible
when they are derived from individuals
of the same species ; it is only then that
they exercise an attractive influence upon
each other and fuse together in the act
of sexual reproduction. The sexual cells
of Mosses and Ferns, apart from all other
considerations, would not unite because
the spermatozoids of Mosses are attracted
to the female organs by sugar, while
those of the Ferns are stimulated by
malic acid. In the case of Phanero-
gams, a mixed union of sexual cells is
likewise prevented by various obstacles
to pollination and fertilisation. Oc-
casionally, however, the sexual cells of
different varieties, species, or even genera
;. 'J4"). A small inflorescence of
have shown themselves able to unite Kalmia glauca the stamens are re- ;

and produce descendants capable of tained in pocket-like depressions of

the corolla.
development. Such a union is termed
HYBRIDISATION, or bastard -formation, and its products HYBRIDS or
BASTARDS.
Through the demonstration of the possibility of hybridisation,
the sexuality of plants, for a long time doubted, was indisputably
proven. With this view, and following the work of
object in
FAIRCHILD, in 1717, were raised in great numbers by
hybrids
KOLREUTER as early as 1761 ( 117 ). It also demonstrated that the
real purpose of sexual union was the combination of the properties
of both parents, for transitional forms are found among hybrids
which in many characteristics resemble the male and in others
the female ancestor (heterodynamic hybrids), or they may show an
equal combination of the characters of both (homodynamic hybrids,
e.g. 9 * Nic. paniculata <$ ).
Nicotiana rustica Less frequently it
happens that the hybrid resembles one ancestor almost exclusively.
In such a case the attributes of the other ancestor remain latent, and
may appear regularly or quite unexpectedly, through atavism (p.

299), in later generations.

The mingling of characters is often complete. Had one species simple leaves
and the other compound, would have leaves more or less cleft or
their hybrid ;

were the flowers of one parent species red and those of the other yellow, tin-
hybrid frequently bore flowers with red and yellow markings (mosaic hybrids), or
314 BOTANY PART i

which were orange-coloured. If an early blooming form were crossed with a

late bloomer, the hybrid would flower at a time intermediate between the two.
A large number of spontaneous hybrids have been found which have arisen
naturally from plants with a special tendency to hybridisation. That such
natural hybrids do not oftener occur is due to the lack of an opportune time or
space for their development, and also to the fact that in the case of pollination
of flowers with different kinds of pollen, that of their own species seems as a
rule more effectual in effecting fertilisation ( 118 ).
The investigations of DE VUIES, CORRKXS, TSCHERMAK, and others, have
greatly increased our knowledge of the laws governing the inheritance of different
characters of plants. No final conclusions are yet attained, however, since even
in regard to limited portions of the problem new features are still becoming
119
apparent ( ).
In the crossing of closely related forms (varieties or nearly related species),
Mendel's laws (called after their first discoverer, GREGOR JOH. MENDEL [1866] are
found to hold. These laws, the most generally applicable of which is that of the
segregration of characters, were independently rediscovered by DE VRIES and
CORRENS. An example will give the best idea of these laws. If a red-flowered
Mirabilis jalapa be crossed with a white-flowered individual one obtains a
generation of hybrids with uniformly rose-coloured flowers. If these are fertilised
from one another a second generation is obtained, but the individuals of this are
not uniformly coloured in addition to rose-coloured plants pure red-flowered
;

and white-flowered plants occur in the proportion per cent of 50 25 25 i.e. : :

in the ratio 2:1:1. When fertilised from one another the pure red-flowered
plants produce a red-flowered progeny and the white-flowered plants also breed
true they have returned to the pure parent forms.
;
The 50% of rose-coloured
plants again splits in the next generation, and like the former generation yields
25% pure red, 25% pure white, and 50% rose-coloured plants. The proportion of
hybrid plants thus continually becomes lessened by the return to the red and
white types in the eighth generation only 0'75% of hybrids remain, and this
;

small remainder continues to split further on breeding. These results are

theoretically explained by assuming that the sexual cells of the rose-flowered
hybrids are not themselves of hybrid nature, but are already segregated into
pure red and pure white sexual elements. In the process of fertilisa-
O~O tion the union producing a hybrid, red x white (white x red, red x white)
y will occur twice as frequently as the union red x red or white x white
which give rise to pure forms. This is illustrated in the accompanying
diagram in which the dark circles represent the red sexual elements.
The characters in which the parents differ do not, however, always blend so
that the hybrid exhibits an intermediate character. More usually the hybrids
completely resemble in this respect either the paternal or maternal parent, the
character of the one parent being dominant in the hybrid while the other
remains latent (Law of. dominant characters). This is the case for example in
hybrids between Urtica pilulifera with serrate leaves and U. Dodarti (Fig. 246).
The hybrids have all serrate leaves like U. piliiliftra, so that in the second
generation the proportion of serrate-leaved to entire-leaved individuals is per cent
75 :25 (3 1). Only 50% of the serrate-leaved individuals are, however, of hybrid
:

nature and continue to show a similar splitting of characters in the next genera-
tion 25% have become pure U. piluUfera.
;
It is impossible to predict which
characters will prevail in any cross, and the question can only be settled by experi-
ment usually the phylogenetically younger character appears to be dominant.
;
 PHYSIOLOGY 315

When the parents differ in two characters instead of only one, instead of
monohybrids, dihybrids result. It then appears that the several characters

are independently transmitted and distributed in the descendants. Thus new

combinations of characters may come about, a fact of great importance in plant-

Fi. 24(5. Hybridisation of Urtiea pilulifera (to the left, above) with U. Dodarti (to the right).
I, First generation which, though hybrid, shows the dominant leaf-form of U. pilulifera. Its
sexual cells, in which a separation of the character occurs, give rise in generation II to a pure
and constant pilnlifi'i-n, apure Dodarti and two hybrids. In generation III the pure forms to
the right and left arc constant, the hybrids in the centre show furthersplitting of characters.

breeding. From the crossing of peas with yellow, wrinkled seeds and those with
green, smooth seeds, among other possible combinations of the characters the new
ones yellow-smooth and green-wrinkled appear (autonomy of characters). For two
pairs of characters the ratio is 9 3 3 : 1, only four externally distinguishable com-
: :

binations resulting, though the nine individuals are really different, the differeuce
being masked in part by the dominance of certain characters. If is the number m
316 BOTANY PART i

of pairs of characters, the number of internally distinct combinations is 3 m and the ,

number of constant forms 2 m If there are four pairs of characters there thus
.

result 81 hybrid-combinations, and among these 16 constant forms.

Many characters, however, tend to remain associated together (coupled
characters).
many races quite new characters frequently appear in the hybrids.
In crossing
Thus in crossing a white-flowered with a yellow-flowered Mirabilis there may result,
besides white-flowered and yellow-flowered descendants, others with rose-, red-,
red-white, yellow-white, rose-red, and rose-white coloured or striped flowers.
Since these newly appearing characters each in turn obey the Mendelian laws it
is assumed that they were latent in the parents, and that the latter belonged to

what are called crpytomerous races.

The Mendeliau laws hold also in the animal kingdom and for the human race,
but only for the crossing of nearly related forms. Thus crosses between negroes
and negro-albinos follow Mendelian laws, while those between negroes and whites
do not.
The more closely allied the parent plants, the more readily, as a rule, may
hybrids between them be produced. Many families seem to incline naturally to
hybridisation (Solanaceae, Caryophyllaceae, Iridaceae, etc.) others again develop
;

hybrids only occasionally or not at all (Papilionaceae, Convolvulaceae, Coniferae,

etc.). Even in the same family the related genera and species exhibit great
differences in the readiness with which they may be crossed. The Grape-vine, and
also the Willow are easily crossed with other species of their own genus, and the
same also true of the different species of Dianthus, while the species of Silene
is

cross with each other only with difficulty. Species hybrids are easily produced
from species of Nieotiana, of Verbascum, and of Geum on the other hand, it is
;

very difficult to cross different species of Solanum, Linaria, or Potcntilla. The

hybridisation, however, of nearly allied iorms is often impossible the Apple with
the Pear, for instance although the Peach and Almond may be crossed, and also
the species of even the different genera Lychnis and Silene, Rhododendron and
Azalea, Aegilops and Triticum, Secalc and Triticum (RiMPAu), Zea and Euchlacna
(Zea canina results from a cross between Z, mais <J and Euchlaena mexicana ? ),
each to their
" sexual
according affinity."
DERIVATIVE HYBRIDS arise when hybrids
are crossed with one another, or with
one of the original parent forms. In this way it has been possible to unite six
species of Willow in one hybrid, and in the case of the Grape-vine even more species
have been combined. It is only in rare cases, however, that the form of the hybrid
remains constant in the succeeding generations. These exhibit more frequently a
tendency to revert to one of the original ancestral forms.

In addition to their inherited qualities HYBRIDS EXHIBIT NEW

PECULIARITIES not derived from their parent forms. These are a
MODIFIED FERTILITY, GREAT TENDENCY TO VARIATION, and often a
MORE LUXURIANT GROWTH. The fertility is often so enfeebled that
the hybrids either do not flower (Rhododendron, EpiloUum), or are sterile
and do not reproduce themselves sexually. This enfeeblement of the
sexuality increases the more remote is the relationship of the ancestral
forms. Other hybrids such as those of Salix and Hieracium remain
fertile.
The tendency to variability is often greatly enhanced in hybrids,
FIG. 247. Laburnum Adami, Poit (Cytisus Adami, Hort) with atavistic branches showing the
characters of the two parental forms, Laburnum, vulgare to the left and Cytisus purpureus
to the right.

317
318 BOTANY PART I

especially in those arising from the hybridisation of different varieties

of the same species.

Hybrids, particularly those from nearly related parents, frequently

produce more vigorous vegetative organs, they bloom earlier, longer,
and more profusely than the uncrossed plants, while at the same
time the flowers are larger, more brilliant, and exhibit a tendency to
become double. The luxuriance of growth and the increased tendency
to produce varieties displayed by the hybrids have made the whole
subject of hybridisation one of great practical as well as theoretical
importance.
It is doubtful if hybrid forms can be produced (graft-hybrids)

by a vegetative union of portions of two different plants (grafting,

budding) ; Laburnum Adami (Fig. 247), and more recently an inter-
mediate form between Mespilus germanica and Crataegus monogyna ( 12 ),
appear to be of this nature. As yet, however, attempts to experi-
mentally obtain these forms by vegetative or sexual methods have
not succeeded. (Cf. p. 252.)

The Dissemination and Germination of Seeds

the seeds after their separation from the parent plant simply
If
fell earth, the young seedlings would be injuriously restricted
upon the
to the place already occupied by the parent plant, and would also

spring up in such large numbers that they would mutually ex-

terminate each other. The dissemination of the seeds thus becomes
a necessity, and although a larger or smaller proportion perish in the
process, a small number eventually find themselves in a favourable
environment. !

For their DISSEMINATION, seeds (and also spores and other repro-
ductive bodies) make use of the same agencies as are employed for
the conveyance of pollen. Thus their dispersion is effected by means
of currents of air and water, by their forcible discharge from their

receptacles and by animals to these means must now be added

;

human intercourse as a by no means unimportant mode of dispersal.

To ensure the dispersal of seeds by the wind, all those contrivances are of use
which serve to increase their with but small augmentation of their
superficial area
weight. Of this nature are the hairy appendages of seeds and fruits, as in
Gossypium, Epilobium, Populus, Salix, Typha, Clematis, and the fruits of the
Compositae with their pappus, of Valeriana, etc. Compared with the accelerated
fall in a vacuum, the retardation exerted
by the resistance of the air (by which the
opportunity for dispersal through the agency of the wind is enhanced) in the case
of Cynaria Scolymus is, in the first second, as six to one. Similar adaptations
for utilising the agency of the wind as a means of dispersal are the wing-like

appendages formed from the expansion of the sepals (Diptcroearjms) or of the

ovary (Acer, Frazinus, Ulmus, Polygonum, Robinia, Gleditschia, and the fruits of
many Umbelliferae), or of the seeds themselves, as in the winged seeds of the
SECT. II PHYSIOLOGY 319

Bignoniaceae and many Terustroemiaceae). In a seed of one of the Bignoniaceae

(Fig. 248), with its widely outspread, glossy wings, the centre of gravity is so
disposed that the seed floats lightly along through the air in an almost horizontal
course, and with a motion like that of a butterfly. The seeds of Zanonia, one of
the Cucurbitaceae, are very similarly equipped. In the Lime the subtending leaf
which is attached to the inflorescence is retained to facilitate the dispersal of the
fruits by the wind and in the seeds of the Fir the winged appendages are
;

derived from the tissue of the placental scale. The aerial transportation of seeds
and fruits, winged only on one side, is accompanied by a continuous spirally
twisting movement which assists to retard their fall. Thus in the above-mentioned
Bignonia DINGLER found the retardation in the first second amounted to thirty
times the free fall, and in Pinus silvestris to seven times.
The diminutive size of many reproductive bodies, and the proportionate
enlargement of their surface in comparison with their volume, increase their
buoyancy. Microscopically small Fungi, spores, and Bacteria are in consequence

FIG. 248. Winged seed of Pithtcocteniuni echinatnnt. (Nat. si/i*.)

easily transported by the wind. According to FALCK'S observations even the heat
produced by the respiration of a fructification of a fungus is sufficient to suspend
the spores in the air. In the spores of Lycopcrdon caelatum DINGLER found the
retardation to be as 1 to 1000, which, according to NAGELI, could only be theoretic-
ally explained by the supposition that the retardation was intensified by a thin
121
layer of air permanently adhering to the surface of the spores ( ).
Minute dust-like spores and seeds form the most effective means of dispersal, as
was shown in the return of vegetation to the island of Krakatoa, where the flora
was destroyed by a volcanic eruption in 1883. Following the forms with minute
seeds or spores came those with hairy fruits or seeds, and later than these the
winged seeds.
Seeds and fruit are also frequently transported great distances by the agency of
VVATI.K. In the case of maritime plants the seeds are often especially adapted
(water-tight floating tissues (Fig. 249) large air-spaces serving as swimming-
;

bladders, etc.) for transport by ocean currents. Through the possession of such
devices, the seeds of West Indian plants are carried to Norway by the Gulf
Stream, as was known to LINNAEUS, and the appearance of Coco-nut palms as the
first vegetation on isolated coral islands is in like manner due to the adaptation of
their fruits to transport by water. SOHIMFEB was able to show in detail the value
of ocean-currents for the geographical distribution of the strand-flora ( 122 ).
ANIMALS participate largely in the dissemination of seeds ;
either by eating the
320 BOTANY

agreeably tasting and often attractively coloured fruit, and excreting tin- un-

digested seeds, or by their involuntary transportation of seeds and fruits which

have become in some way attached to them. This is effected in many cases by
hooks and bristles (Lnppa, Galium aparinc, B-ulciix, 7:>A ///"*/;<//// //w. Xiniiliinm,
and the fruits <>f M-il-icago minima, so common in sheep's wool and erroneously
termed wool-lice). The seeds may also become attached to animals by means of
some sticky substance in this way the seeds of the Mistletoe, which stick to the
;

1
leaks of birds eating the berries, finally adhere to the
brandies of trees upon which the birds wipe their
bills. The widespread distribution of fresh-water
plants can only be accounted for through the agency
aquatic birds.
<>!'

The natural distribution of plants has been greatly

modified by the interference of man, especially in these
days of universal commercial intercourse by rail and
sea. By their instrumentality not only have the
useful plants been widely distributed over the earth,
but the weeds have followed in the same way and ;

many a seed thus accidentally carried to other lands

has finally found there a new place of growth.
The forcible discharge of spores and seeds is effected
by the sudden liberation of hygroscopic or tissue
tensions. It has already been mentioned that the

eapillitia of the Myxomycetes and the elaters of

the Liverworts serve for the dispersal of the spores.
In the case of the Box (Bumis), the smooth seeds
are forcibly discharged by the contraction of the
pericarp, like a bean pressed between the fingers.
The dry fruit of Hura crepitans bursts apart with a
report like that of a pistol, and is scattered in pieces
Flo. 249. Fruit of Termiiialia far and wide. The turgescence and elasticity of the
cctiappa from the drift. In cell-walls give rise to the tension which results in the
tto upper ^ecimea the corky
forcible abjection of the sporangia of Pilobolus (Fig.
floating tissue is seen from the
outside, the smooth, super-
224 )> and m . ,.
the ejection ot the ascospores ot many
ticial layer being already worn Ascomycetes. The bursting and rolling up of the
The lower
away. figure is
segments of the seed-vessels of Liipaticiin, by means
of a cross section through of which the <ji spersa i O f t h e 8e eds is effected, are due
the floating tissue ; the seol , . . - . . _,. ,. .

has been removed to t ' ie su dden release of tissue-tensions. The fruits

(\fter
S( JIIMPER.) of Momordica elaterium and Ecballium have the
pericarp distended by the pulpy mass within when ;

the mature fruit becomes detached from the stalk which fitted like a champagne
cork into its place the fluid contents with the seeds are forcibly squirted from
the opening.

Germination. The first condition for successful germination is

that the seed should retain its vitality during its dispersal. The
small amount of water contained in the seed and the arrest of the
main processes of life render the resting seed resistant both to ex-
tremes of temperature and to drying. Firm investments serve to
protect it from mechanical injury. The second condition of germina-
tion is that the seed should reach suitable soil, and this is effected by
 PHYSIOLOGY 321

the assistance of wind, raiu, and animals aided by the existence of

cracks and crevices in the ground. The fixation of the seed in the
soil, which is often of importance for the emergence of the seedling
from the testa, is assisted by structural peculiarities of the surface
such as furrows, ridges, bristles, and
hairs ; the latter may be able to expand
hygroscopically.

Tin- fruits of the Geraniaceae (Erodiurii,

Fig. 222) and Gramineae
(Stipu, Aveim utmlix.
and species of ^Ir-isfida) are enabled, by means
of movements due to hygroscopic torsion as-
sisted by the presence of stiff backwardly-

directed bristles to bury themselves in tin-

ground. In the cast- of Ti-lj'nl In-lit, subter-
runcuni and v//v/'7//.s- hypogtua&e same result
is accomplished by the geotropic growth of the
fruit-stalks, while the seed-capsules ofLinariu
cymbaJaria are deposited in the crevices of
walls and cliffs by the negative heliotropic
movements of the fruit-stalks. Nuts, acorns,
and seeds buried by squirrels or other animals
in the ground and forgotten, or for any reason
not made use of, often germinate. The seed-
lings of Mangrove trees, llhr.nplwru, Bruguiera,
and Kandclia (Fig. 250), exhibit a most
peculiarmanner of growth to ensure their
lodgment in the ground. The seed germinates
in the fruit before it is detached from the tree.
When the radicle, has attained a considerable
length, the young seedling, separating either
from the cotyledons or from the fruit-stalk,
falls to the earth, boring like an arrow into
the mud, in which it sticks erect and is thus
enabled to commence its growth without del.-i\.
In some species of the Mimosaceous genus
Inya naked embryos are also liberated from tin-

fruits or seeds.
I'n.. -'.Mi. Kinnlelia llAeedii. The mass-
Manyseeds and fruits acquire a more or
ive root the seedling (/) has broken
(it
lessvoluminous MUCILAGINOUS SHEATH, which out of the fruit. When the ]>l;mt
serves a double purpose. Quince seeds, Flax separates from the fruit the root will
seeds, seeds of the Plantain, of Crucifers, tin- become inserted into tin; muddy soil.
fruits of S" //-in Imi-inimiiii, seed of Cuphea and (After S( HIMPER'S /V,i,,M,Vo,,n</>/ry.)

CobfK'u (in the

mucilage which delicate
cells of

thickening bands are rolled up), afford the best-known examples of such slimy
envelopes, which, in addition to fixing the seed to the ground, serve to absorb
water by holding it in their substance or drawing it in hygroscopically (cf.
Mistletoe berries). Fruit -walls, by their spongy nature, may also serve a>
water-carriers (ripe fruits of Tropaeolum, Pvti-i'ii' m xfi'tnosum, Medieuyo terebellmn).

In the soil the seed swells quickly or more slowly and then at
Y
322 BOTANY PART i

once or after a resting period (dependent in part on delay in swelling

due to hard seed coats, partly on the time needed for the growth of
the embryo) commences to germinate. In some cases the germina-
tion (both of seeds and of fungal spores or pollen grains) is dependent
on the presence of special substances or of the host plants (Orobanche,
Tozzia, spores of Bulgaria) in other cases of light (Viscum, Drosera,
Pitcairnia mdidifolia, Veronica peregrina, Nicotiana). In the case of
fern-spores the action of light can be replaced by warmth, and in
moss spores by the presence of sugar in the solution. The action
of light is on the other hand inhibitory in the germination of
123
Acanthostachys strobilacea and Phacelia tanacetifolia ( ).

The seeds of many Conifers do not germinate for several years ; those of the
Ash and Hornbeam in 2 and those of Euphorbia cyparissias in 4-7 years. Some
plants again, in addition to seeds which germinate in the first year, produce others
which require a longer rest (Trifolium pratense, Robinia pseudacacia, Cytisus
Laburnum, Reseda Euphorbia exiyua, species of Dianthus, etc.).
lutea, Even
under favourable circumstances such seeds do not germinate until a definite
length of time has elapsed. Treatment with strong sulphuric acid may in such
cases increase the permeability of the seed-coat and hasten germination. Germina-
tion may be delayed also by external conditions, and the vitality of the seed may
still be retained for years. Thus, for example, on the removal of a forest from land
that had been under cultivation for forty-six years, PETER found that a great
variety of field-plants at once sprang up as soon as the requirements for their
germination were restored. The plants developed under such circumstances are,
124
however, often weakly ( ).

Germination, according to the observations of KLEBS, is intro-

duced by true processes of growth, which result in THE RUPTURE OF
THE SEED COVERINGS ( 125). This is effected either
by the growing radicle, or, in many Monocoty-
ledons, by the cotyledon.In other seeds enclosed
within a shell, the bursting of the latter by
the growth of the endosperm or cotyledons pre-
cedes germination. In cases where the shell is
very hard and does not consist of two halves
easily separable by internal pressure (as in
Walnuts and Cherry-stones), special places are
.
tl . often xprovided for the egress of the voung
FIG. 251. Section through
the upper part of the seedling,
fruit of A crocomia sdero-

S, The hard g ucn and hard

carpa.
pl aces are often present in the thick
S hell;P, the plug which
is pushed out of the
seed . coats of m
r lms* At the end of a Coco . nut for
.

shell by the genninat- example, three points of egress, behind the thinnest of
ing embryo, K;E, endo- which the embryo will be found embedded in the endo-
sperm. (After PFITZER.) sperm, are very easily seen. Through the extremely hard,
thick shell of another Coco-palm, Cocos lapidea, there
are three long germinal pores, while the seedling of Acrocomia sclerocarpa has

only to push ti loosely fastened plug out of the thick shell of the seed (Fig.
SECT. II PHYSIOLOGY 323

126
251) ( ). Similar contrivances are found in the case of Pandanus, Canna,
Typha, Potamoycton, and many Dicotyledons J Tetragonia expansa, Medicago,
and some species of Onobrychis and Portulaca).

SEEDLINGS PENETRATE THE SOIL by means of the elongation of

the primary root, or of the hypocotyl, or also, as is the case with
many Monocotyledons, through the movements of the geotropic
cotyledons. After the descending part is firmly attached to the
soil, by either root- hairs or lateral roots, THE UPWARD GROWTH
COMMENCES. In this process the cotyledons may either remain
within the seed (hypogeal) or unfold above ground (epigeal). The
first is often the case where the cotyledons are full of reserve
material (Phaseolus multiflorus, Aesculus, Quercus), or where their
function is to absorb nourishment from the endosperm (in Palms and
the scutellum of Grramineae). More frequently the cotyledons are
pushed above ground, and may then be thick and filled with reserve
nourishment, or thin and turning green on exposure to the light
In many Monocotyledons, as also in Eicinus, etc., the cotyledons,
even if they afterwards appear above ground, may first take up the
nutritive substances of the endosperm while in the Conifers the
cotyledons perform the same office above ground.
THE COTYLEDONS ARE DRAWN FROM THE SEED by the curvature
of the hypocotyl or of the petioles of the cotyledons (Smyrnium,
Delphinium). The seed-coverings also are often further ruptured by
the swelling of the hypocotyl (Cucurbitaceae, Fig. 231, etc.).
The unfolding of the first leaves above ground is frequently
accompanied by a CONTRACTION OF THE ROOT. This is caused by
the cortical cells of the root becoming more stretched in a transverse
than a longitudinal direction by their turgescence. In consequence
of this the root becomes thicker and shortens considerably. The
seedling is in consequence drawn deeper into the soil, and its position
rendered more secure. Even older plants, particularly those whose
leaves form a radical rosette, notwithstanding their upward growth,
are held close to the ground through a similar contraction of their
roots. The shortening is indicated by the transverse wrinkling of
the surface of the root. According to the researches of RIMBACH the
shortening amounts to 30 per cent of the length of root in Allium
ursinum, to 50 per cent in Arum maculatum, and even to 70 per cent
12:
in Oxalis elegans ( ).

When its attachment in the soil is properly provided for, and

after the first germ-leaves are unfolded, the young plant has acquired
the capacity for self-sustenance, its further growth and development
being dependent upon its own activity.
 PART II

SPECIAL' BOTANY
 SPECIAL BOTANY
SPECIAL BOTANY concerned with the special morphology and
is

physiology of plants. While it is the province of general botany to

investigate the structure and vital processes of the whole vegetable
kingdom, it is the task of special botany to interpret the structure
and vital processes of its separate divisions. The aim of general
morphology is to determine the phylogenetic derivation of the
external and internal segmentation of plants, and to refer their
numerous structural peculiarities to the primitive form from which
they have arisen. The purpose of special morphology, on the other
hand, is to trace the development which has been reached in the
different divisions of the plant kingdom, to understand the form of
individual plants, and to trace the connection between one form and
another. Thus the methods of special morphology are also phylo-
genetic, and furnish the basis for a NATURAL SYSTEM of classification
of the vegetable organisms, based upon their actual relationships.
Although such a system must necessarily be very imperfect, as it is
not possible to determine, directly and indisputably, the phylogenetic
connection of different plants, but only to derive indirectly their
relationships from morphological comparisons, the aim which we set
before us is none the less both legitimate and essentially justified.
Such a natural system, founded on the actual relationship existing
between different plants, stands in direct opposition to the ARTIFICIAL
SYSTEM, to which has never been attributed more than a practical
value in grouping the plants in such a manner that they could easily
be determined and classified. Of all the earlier artificial systems,
the sexual system proposed by LINN^US in the year 1735 is the
only one which need be considered.
LINN.&TJS, in establishing his classification, utilised characteristics which referred
exclusively to the sexual organs, and on this basis distinguished twenty-four classes
of plants. In the last or twenty- fourth class he included all such plants as were
devoid of any visible sexual organs, and termed them collectively CRYPTOUAMS.
Of the Cryptogams there were at that time but comparatively few forms knowin
and the complicated methods of reproduction of this now large class were absolutely
unknown. In contrast to the Cryptogams, the other twenty-three classes were
distinguished as PHANEROGAMS or plantfe whose flowers with their sexual organs
327
328 BOTANY I-AKT n

could be easily seen. LINN.EUS divided the Phanerogams, according to the M-XU.I!
character of their flowers, into such as possessed hermaphrodite flowers (Classes
I.
-XX.), and those in which the flowers were unisexual (XXI. -XXIII.). Plants
with hermaphrodite flowers he again divided into three groups those with free
:

stamens (I. -XV.), which he farther distinguished according to the number, mode
of insertion, and relative length of the stamens ;
those with stamens united with
each other (XVI. -XIX. ); and those in which the stamens were united with the
pistil (XX.). Each of the twenty -four classes was similarly subdivided into
orders. While some of the classes and orders thus constituted represent naturally
related groups, although by the method of their arrangement in the artificial
system they are isolated and widely removed from their proper position, they
include, for the most part, plants which phylogenetically are very far apurt.

LINNAEUS himself (1738) felt the necessity of establishing natural

families in which the plants should be arranged according to their
"
relationships." So long, however, as the belief in the immutability
of species prevailed, the adoption of a system of classification ex-
pressive of relationship and family could have no more than a hypo-
thetical meaning, and merely indicated a supposed agreement between
plants having similar external forms. A true basis for a natural
system of classification of organisms was first afforded by the theory
of evolution.
The system adopted as the basis of the following description and
systematic arrangement of plants is the natural system of ALEXANDKR
BRAUN, as modified and further perfected by EICHLER, ENGLER,
and others.

According to this system, we have to distinguish between CRYPTO-

GAMS as the lower division, and PHANEROGAMS as the higher division
of the plant kingdom.

SECTION I

CRYPTOGAMS
The Cryptogams include an extraordinary variety of the most
different plant forms, ranging from unicellular organisms to plants
exhibiting segmentation into stem, leaf, and root. The Cryptogams,
however, are collectively distinguished from Phanerogams by the
mode of their dissemination by SPORES, in contrast to that of the
Phanerogams, which is effected by SEEDS spores are formed also by
;

Phanerogams, but they are not the immediate cause of the origin
and development of new individuals. Seeds are multicellular bodies,
within, which is included the multicellular rudiment or EMBRYO of a
plant; while spores, which, in the case of the Cryptogams, become
separated from the mother plant, and give rise to a new and
 CRYPTOGAMS 329

independent organism, are unicellular structures. Cryptogams may

therefore be termed SPORE PLANTS or Sporophytes, and Phanerogams
SEKD PLANTS or Spermaphytes although previous usage and custom
;

would recommend adherence to the older terms.

The Cryptogams are divided into the following main groups :

I. The THALLOPHYTA,
embracing a great variety of plants whose
vegetative portion may consist of one or many cells in the form of a
more or less branched thallus. Reproduction is both sexual and
asexual, but there is
usually no definite succession of the two modes
of reproduction.
II. The ARCHECONIATAE exhibit a regular alternation of two
generations in their life -history. The asexual generation forms
spores, and is called the From the spore the sexual
SPOROPHYTE.
generation or GAMETOPHYTE
develops ; this bears sexual organs of
characteristic construction, the male organs being called antheridia,
and the female organs archegonia. From the egg-cell contained in
the latter, after fertilisation, the sporophyte again arises. The
Archegoniatae are divided into
1. The BRYOPHYTA, which include forms with a leaf-like thallus,

;is well as
cormophytic forms, with evident segmentation into stems
and leaves. The Bryophytes possess no true roots, and their con-
ducting bundles, when present, are of the simplest structure. The
sporophyte is a stalked or unstalked capsule, which lives semi-
parasitically on the sexual plant.
2. The PTERIDOPHYTA have small thalloid gametophytes the ;

sporophytes exhibit a segmentation into stems, leaves, and roots, and

also possess true vascular bundles ; they thus resemble the Phanero-

gams in structure.

The Thallopliytes and Bryophytes are also characterised as Cellular Plants, in

contrast to the Pteridophytes or Vascular Cryptogams, which, together with the
Phanerogams, are collectively designated Yascular Plants. Bryophyta and Pteri-
dophyta must be regarded as having had a common origin from the higher Thallo-
phytes, the development of the two groups having been on dill'civnt lines.

THALLOPHYTA
It
A I.

was formerly customary to divide the Thallophyta into Algae,

.Fungi, and Lichens. The Algae are Thallophytes which possess
chromatophores with colouring pigments, particularly chlorophyll ;

they are, therefore, capable of assimilating and providing inde-

pendently for their own nutrition. The Fungi, on the other hand,
are colourless and have a saprophytic or parasitic mode of life.
Such a method of classification, however, although possessing a
physiological value, has no phylogenetic significance, as it does not
express the natural relationships between the various groups. In
330 BOTANY PART n

the Lichens (Lichenes), which were formerly regarded as simple

organisms, the thallus affords an instance of a symbiosis of Algae
and Fungi. From a strictly systematic standpoint, the Fungi and
Algae composing the Lichens should be classified separately, each in
their own class ; but the Lichens, among themselves, exhibit such a

similarity in structure and mode of life, that a better conception of

their characteristic peculiarities is obtained by their treatment as a
distinct class.
The phylogenjetic connections of the fourteen classes into which
the Thallophyta are divided are expressed, so far as is possible, in
the following scheme :

^Bacteria, Bacteria.
\Cyanophyceae, Blue-green Algae.
m
3Ii/.mmycetes, Slime-Fungi.
'Pervlineae, Dinoflagellates.
Conjugates, Conjugates.
0> : Diatomeae, Diatoms.
rf Heterocontae.
s *<' 1/1 1
trophy ceae,Green Algae.
.gf tCJiaraceae, Stoneworts.
3 Rhodophyceae, Red Algae.
^}
!5
S Eumycetes, Fungi.
^ Phycomycetes, Algal Fungi.
Phaeophyceae, Brown Algae.

The Bacteria and Cyanophyceae are among the most simply organised Thallo-

phyta ;they are closely connected and are often grouped together as the Schizo-
phyta. They occupy an isolated position in contrast to the remaining simple-
Thallophytes, which with greater or less probability may be derived from the
Flagellatae. The Flagellatae used to be (and frequently still are) placed with the
lowest animals. As a matter of fact they combine plant and animal characteristics,
and may also be regarded as the starting-point of the lower animals. The
Myxomycetes may also have sprung from them as a group of colourless saprophytes.
The Peridineae are a further-developed branch of the Flagellatae. The simplest
forms among the Heterocontae, the Green Algae, and the Phaeophyceae connect
directly with the Flagellata on the other hand a direct connection of the latter
;

with the Conjugatae and Diatomeae (which together form the Zygophyceae), while
probable, is not shown in existing forms.
The Phycomycetes have. branched off from the main series of the Chlorophyceae.
The origin'of the Red Algae and the Eumycetes, which appear to have sprung from
a common stock, is still in doubt. The Characeae occupy a quite isolated and very
advanced position, and are usually regarded as the most highly developed of the
Green Algae.

The Thallophytes are commonly multiplied and distributed by

asextially produced spores, the mode of development of which differs
in the several groups. In many cases the spores arise by a process
SECT, i CRYPTOGAMS 331

of cell division within certain cells, which are known as sporangia ;

in other cases they arise by modification and separation of cells of
the thallus or by a process of cell-budding. When the spores possess
cilia and are able to move actively in the water,
they are known as
swarm-spores (zoospores) ; when they do not bear cilia they are
termed aplanospores. In the latter case the spores if distributed
by water may be naked, or they may be provided with a cell-wall
and suited for distribution in the air.
Sexualreproduction is also of wide-spread occurrence. It
consists, the simplest cases, in the production of a single cell,
in
the ZYGOSPORE or ZYGOTE, by the union or conjugation of two
similarly formed sexual cells or gametes. The organs in which
the gametes are formed are termed gametangia ; planogametes
are provided with cilia while aplanogametes are non-ciliated. In
many of the more highly developed forms, however, the gametes are
differentiated as small, usually ciliated, male cells or SPERMATOZOIDS,
and as larger non-ciliated female cells, the egg-cells or OOSPHERES.
The spermatozoids are formed in antheridia, the oospheres in oogonia.
As a result of the fusion of an egg-cell and a spermatozoid, an
OOSPORE is The first form of sexual reproduction or
produced.
fertilisation termed ISOGAMOUS, the second OOGAMOUS ; but these
is

are connected by intermediate forms. It must be assumed that the

sexual cells have been derived in the phylogeny of plants from
asexual spores, and that asexual multiplication has taken origin from
simple cell division. The gametangia, oogonia, antheridia, and
sporangia of the Thallophyta are homologous structures. The sexual
reproduction has originated independently in several distinct groups.
While the reproduction of some Thallophyta is exclusively asexual, and of others
exclusively sexual, in many others both forms of reproduction occur. In the latter
case this may occur on the one plant, or separate successive generations may be
distinguishable. Generally speaking, there is, however, no regular succession of
asexual and sexual generations in Thallophytes, the mode of reproduction being to
a great extent under the influence of external conditions ( 1 ). Only in some Brown
Sea- weeds, in theRed Sea-weeds, and some Fungi is there an alternation of a
sexual generation (gametophyte) with an asexual (sporophyte), such as is found
in all Bryophytes and Pteridophytes.

CLASS I

2
Bacteria ( )

Bacteria are unicellular or filamentous organisms of very simple

construction. Chlorophyll is wanting in them, and their mode of
life is consequently a parasitic or
saprophytic one. A
large number
of species exist distributed over the whole earth, in water, in the
3.32 I IOTA NY r.uir n

soil, dead or living plants and

in the atmosphere, or in the bodies of
animals. They are often termed Fission-Fungi, or Schizomycetes,
since the multiplication of the unicellular forms takes place by a
division into two and the separation of the segments. This mode
of multiplication is also found in other unicellular plants.
The cells of the Bacteria arc surrounded by a thin membrane, and
contain a protoplasmic body, which is usually colourless, and can
be made to contract away from the membrane by plasmolysis.
The protoplasm may contain one or more vacuoles. One or several
granular structures are also present in the protoplast; these so-called
chromatin bodies may be deeply coloured by stains, and have been
regarded as nuclei by various authors. Since, as yet, undoubted karyo-
kinetic division has not been observed in these bodies, the presence of
nuclei in the bacterial cell cannot be regarded as certainly established.
For the most part the Bacteria are extraordinarily minute organ-
isms, and probably include the smallest known living beings. The
spherical cells of the smallest forms are only O'OOOS mm. in
diameter ; the rod-shaped cells of the tubercle bacillus are only
0'0015-0'004 mm. long, while the transverse diameter of most
speciesis about O'OOl mm.

The simplest form of Fission-Fungi are represented by minute

spherical cells, COCCI. Forms consisting of rod-shaped cells are
designated BACTERIUM or BACILLUS. Rod-shaped forms with a slight
spiral curvature are called VIBRIO, and those more strongly curved
SPIRILLUM. Straight filamentous forms are termed LKPTOTIIJ:IX,
spirally wound filaments, SPIROCHAETE. In the highest stage of their
development the Fission-Fungi consist of cell filaments exhibiting
false branching. The unicellular cocci, rod -shaped forms, and
vibrios may also remain united in chains after the cell-division.
Frequently the cell -membranes undergo a mucilaginous swelling,
the cells or cell-rows being embedded in the gelatinous mass. This
stage of development is termed ZOOGLOEA.
Many Bacteria are motile. Their independent movements arc due
to the vibration and contraction of fine protoplasmic cilia. These
according to A. FISCHER,
flagella, are either distributed over the whole
surface of the cells (peritrichous)
(e.g.
Bacillus subtilis, Fig. 254 a, d;
Bacillus typlii, c; Bacillus tetani, Fig. 257 e), or they spring
Fig. 252
from a single point either as a single flagellum (monotrichous) or
as a group (lophotrichous). A
single, polar flagellum occurs
in
niirio cholerae (Fig. 252 a) ; a polar terminal tuft of flagella in Spiril-
lum a in/ ula (Fig. 252 b, d) ; a lateral tuft in the swarm-spores of
Cladotlirix (Fig. 253). The ciliary tufts may become so closely inter-
twined as to present the appearance of a single thick flagellum, The
cilia are never drawn within the body of the cell, but undergo
dissolution before the formation of spores takes place, or under
unfavourable conditions (Fig. 252 e).
 CRYPTOGAMS 333

Multiplication of the individual is accomplished vegetatively by

the active division or fission of the cells the preservation and dis- ;

l-'h.. _' ">_'.

Types c.l' arrangement of Hagclla. .

1'ihrin I'lo'li,'" ;
n, il, fi-]iii'!/ln HI
n mini" ; il,

development of a new bunch of <-ilia in divi- Fio. 253. Cladothrix dldwtoma. Formation
sion ; ', Ihicilflix l/li'lti', i. liiirillli* siilitili*. of swarm cells from the cells of the lila-

(After A. FISCIIKK, X 22">0.) nient. (After A. FISI-HKK, x 1000.)

tribution of the species by the asexual formation of resting spores.

These arise as endospores (Figs. 254 c, 256 e, /) in the middle or
at one end of a cell by the inner portion of the protoplasm separat-
ing itself from the peripheral, ,

and surrounding itself with

a thick membrane. The
membrane of the mother cell
becomes swollen and disin-
tegrated when the spore is

ripe. Spores are not found in

all species.

Bacillus sitbtilis, the Hay bacillus

(r'ig. 25-i), which appears as a rule
in the decoction obtained by boiling
hay in water, will afford an example
of the life-history of a Bacterium.
The spores of this species, which
withstand tin- clfect of the boiling

water, produce on germination rod-

shaped swarming cells with cilia
on all sides these divide and may
;

remain connected in short chains.

At the surface of the fluid these
f'n.. -'.'.4. / a, <l, Motile cells and
swarming cells change into non- chain of cells; /;,non-motile cells and chains of
motile cells without cilia, which cells: ., -spores from the zoogloea, >. (From A.
divide up, giving rise to long in- FISCHKK. r.j/7. ii lii i- \\dct er'n- ii. n-il, x 1500; <,

tertwined chains of cells. X 250.)

These
are associated together in tlie
pellicle covering the surface (zoogloea stage). Spore
formation occurs when the nutritive substances in the fluid are exhausted.
Although t lie
cycle of forms passed through in the lite-history of a liacterium
334 BOTANY

is a very simple one. the individual species, which can often be barely dis-
tinguished by morphological characters, show great variety in their metabolic
Miid in their mode of nutrition. The majority of Bacteria require
oxygen for their respiration, and are
therefore aerobic; many
however, can,
develop without this gas, while some
species, e.g. the butyric acid bacterium and
the tetanus bacillus, are strictly anaerobic
and only succeed in the absence of
oxygen (cf. p. 242).
Saprophytic and parasitic species are
distinguished, though a sharp SI-JIM rat ion
is often impossible. In cultures the para-
sitic forms can be made to lead a sapro-

phytic life on suitable substrata.

To the saprophytic Bacteria belong in
the first place the forms which inhabit
Fio.. 255.. A, Iso--
IiucoiMMtoe nusenterioide*.. , .,
, ,. ., , ,
/-, , ,

feted without
water The Wldel
"
y distributed Clado-
.,-lls sheath;
gefetbuma
tkritS dichotoma is morphologically the
B, C, formation of chain of cells with
gelatinous sheath D, portion of mature
;
highest among these. It is found in
zoogloea K, formation of isolated cells in
;
stagnant water, and consists of faldv
the filaments ofthezoogloea. (After VAN
branchi delicate filaments attached to
Hi:.. HEM, X 520.)
Algae, stones, and woodwork, and forming
a slimy coating over them ;
the filaments are composed of rod-shaped cells.
Reproduction is effected by ciliated swarm-cells, which originate by division
from cells of the filament and are -set free by the swelling of the sheath (Fig.
253). The swarm-cells come to rest after a time and grow into new filaments.

Fio. 256. Bacteria of fermentation, u-c, Vinegar bacteria ; ,

LV /'<" . /'<>.*.'

c, Bac. Kutzin guinus; d, Bac. addi lactici, lactic acid bacillus ; e, Clostriiiimn imtiti-ii-iim, butyric
acid bacillus;/, Plectridium paludogum, fermentation bacterium from marsh \vutt-r. (I-'miii
A. FISCHER, Varies, uber Bacterien, x 1000.)

Another very common form is Crenothrix Kuhniana, which consists of un-

branched filaments attached to the substratum, but easily broken. It often
forms masses in the cavities of water-pipes, blocking them up and rendering tin-
water undrinkable. The reproduction of Crenothrix is effected by small, round,
SECT. I CRYPTOGAMS 335

non-motile cells, which arise by subdivision of the cells of a filament enclosed by

its sheath.
The numerous kinds of Sulphur Bacteria, of which Beggiatoa alba is the
most widely distributed, are found in sulphurous springs and at the bottom of
pools where sulphuretted hydrogen is being formed by decomposition of organic
material. These Bacteria oxidise sulphuretted hydrogen into sulphur, and store
the latter substance in the form of rounded granules within their cells (cf. p. 243).
Leptothrix ochracea, the so-called Iron -Bacterium, oxidises oxide of iron to
the hydrated oxide of iron which it accumulates in the sheaths of its filaments
(cf. p. 243). It occurs in ditches and swampy places in meadows.
The zymogenous or fermentation Bacteria and the saprogenous or decomposi-
tion Bacteria are other saprophytic forms. The former oxidise or ferment carbo-
hydrates. The decompose nitrogenous animal or vegetable substances
latter
(albumen, meat, with the liberation of ill-smelling gases.
etc.)
Thus Leuconostoc mesenterioides (Fig. 255) causes fermentation of beet-sugar.

FIG. 257. Pathogenic Bacteria, u, Pus cocci h, erysipelas cocci c, gonorrhoea cocci (/, splenic
; ; ;

fever bacilli e, tetanus bacilli

; /, diphtheria bacilli
; g, tubercle bacilli
; h, typhoid bacilli
; I, ;

colon bacilli fc, cholera bacilli. (Prom A. FISCHER, Fortes, uber Bocterien, x about 1500.)
;

It forms large mucilaginous masses like frog-spawn, the bead-like rows of cells

being surrounded by a gelatinous investment. The acetic acid bacteria (Fig. 256
a, b, c) oxidise alcohol to acetic acid. The transformation of sugar into lactic
acid is brought about by the rod-like cells of Bacillus acidi lactici (Fig. 256 d).
Clostridium butyricum (Fig. 256 e) forms butyric acid from various carbohydrates
in the absence of oxygen, while certain marsh Bacteria (Fig. 256 /) in the absence
of oxygen form marsh-gas from cellulose. Bacillus vulgaris is the most common
cause of decomposition of meat, albumen, etc.
The photogenic bacteria produce within their cells a substance which becomes
phosphorescent on oxidation (cf. p. 246). The most widely spread of these phos-
59
phorescent bacteria is Bacterium phosphoreum, and occurs on meat C ).

The parasitic bacteria inhabit both animals and plants. The best known
forms which cause diseases of plants (bacterioses) are Pseudomonas Hyacinthi,
Bacillus phytophthorus, which attacks the potato, and Bacillus Oleae, which gives
3
rise to the Canker of the Olive Tree ( ).
The numerous pathogenic Bacteria are the most important causes of infectious
diseases. Their injurious influence on the tissues and blood of men and animals
is brought about by the excretion of poisonous substances, to which the name
336 BOTANY I-AKT n

toxins has been given. The following forms may be mentioned. ,s

/"y</<///..i-<'.v//>

j>!/iiif<"Hc$ (Fig. 257 a), the cocci of which form irregular or racemose masses, is
tin- most common cause of suppuration, while Xt irptwoccus pyogtncs (Fig. 257 b),
with cocci united in chains, occurs in erysipelas and other suppurative lesions.
.Mi'-rococcus (Z>t^A-<i<ri/N) <ii>norrJivrae (Figs. 257 c, 258 ), has somewhat flalti-n.-ii

cocci arranged in pairs, and causes gonorrhoea. Bacillus n nth rue I a (Figs. 257 d,
258 c) was found by R. KOCH in the blood and organs of animals suffering from
splenic The relatively large rod-shaped cells may be united in short
fever.
chain* they form endospores in cultures in the same way as the Hay bacillus.
:

Hurillus tetani (Fig. 257 e) occurs in the soil, and is the cause of tetanus. Its
straight rod-shaped cells arc ciliated, and grow only in the wound itself; their

>

\\*

Fi<;. li.vs. stained i>vri>aiatioiui from Xie^k-r's Trj(-bovk <>j l'(<th^>njy. a, jjonococci hi the
jronorrha-al discharge, mucus and pus corpuscles with cocci (methylenc Mm- and eositi), x 700:
ft, tubercle bacilli in sputum of plitjiisis (fuclisin and mothylene blue), x 400; c, splenic fever
bacilli in the pustule of tlio disease (methylene blue and vesuvin) x 350. (From A. l-'is. in i:.

Vorlfs. "'"' />'"' ''

r'V.)

spores are formed in the swollen end. Bacillus injtucnzae, short, slender rods :

Bacillus -jifKds. small, stout, non-motile rods. LOFFLER'S Bacillus d iplit/n i-in,
(Fig. 257 /) consists of small rod-shaped cells sometimes thickened at one end.
KOCH'S Bacillus tuberculosis (Figs. 257 a, 258 b), which is found in all tuberculous
lesions and secretions, as in the sputum, is a slender, slightly curved rod. Typhoid
fever is caused by the ciliated cells of Bacillus typhi (Fig. 257 ft) Bacillus coli ;

(Fig. 257 i),the colon bacillus, which is as a rule harmless and always occurs in
the human intestine, closely resembles the typhoid bacillus. The comma bacillus
of Asiatic cholera, Vibrio cholerae (Fig. 257 k), was discovered by K. KOCH. It
occurs in the intestine as short curved rods with a single polar flagellum, and
sometimes in longer chains of spirally wound cells. Spirocliacte obcrmeicri, which
consists nf long, thin, spirally wound filaments, without Hagella, but capable of
motion, occurs in the blood during the attacks of relapsing fever, of which it is
the cause.
Besides the above injurious parasites there are others which are more or less
harmless occurring on the mucous membranes, in the mouth (Fig. 4), or the
intestine. Sarciim ri'iitricvli, which occurs as packets of cocci in the stomach and
intestine of man. will serve as an example of these.
 CRYPTOGAMS 337

Bacteria included under tin- name Bacillus rndiricla ( JUtizobium

Tin:

rinosarum) occur in the root-tubercles of Leguminosae, and, like certain

other Bacteria found in the soil (Azotobacter chroococcum), and in the sea, are
capable of utilising free nitrogen in their metabolism (p. 232). The denitrifying
Bacteria which occur in the soil and in the sea decompose nitrates and nitrite*,
and liberate free nitrogen.
In addition to saprophytic and parasitic Bacteria, there are some which, though
possessing no chlorophyll, obtain their food from inorganic compounds only.
These are the Nitrite Bacteria (Nitrosomonas), and the Xitrate Bacteria (Nifi--
lnn:ti-r which live in tin- soil.
, The
former oxidises ammonia to nitrous acid,
and the latter oxidises tin- nitrous to
nitric acid. They both obtain their
carbon from carbonic acid, and thus
derive their food independently of any
organic food-supply (Fig. 259, cf. p.217).
The Myxobacteriaceae (
4
) are a very
peculiar family of Bacteria, our accurate
knowledge of which is in the first instance
due to THAXTKI:. They nearly all live
saprophytically on the dung of animals,
and in habit resemble the My.\omycetes
(cf. p. 340). In the vegetative stage they
appear as swarms of rod-shaped Bacteria
connected together by the gelatinous sub-
stance secreted by the cells, and exhibit

-
FK;. J00. A, Myxueoeeus iligitatuy, bright
ivd tVuctilicatinn occurring on dung (x
120). /;, I'nlycnyiuiii jrriinigenium, red
fructification on dog's dung (x 40). C,
Fiu. li'.'.i.
Nitrifying laeteria. alter Wiiidgrad- L'hniiiii-oiiiyres fy>iVi</f orange fructifica-
i/.<,

sky. a, Nitrotomonuewropeta, from Zurich ; tion on

anteloi>e'8 dung. D, young fructi-
li, Xili-osoiiniiiiix jin-intfiixi?, IVoni .Java ; <, fication (x 45). /-'. single cyst germinat-

Xitriibt'fti r, from Queto. (From Kis< HI i:.

ing (x -'00). (.1. U. after gi'KiiL : ''/:,
Varies, -iiber liacterien, x 1000.) after THAXTF.R.)

slow creeping movements. Ultimately they form fructifications that are

usually
brightly coloured these have the form either of definitely limited masses of spores
;

or of cysts containing within a firm membrane the numerous The cysts

spores.
are unstalked, or are raised singly or in groups on a stalk, formed, like the wall of
the cyst, of the hardened gelatinous material (Fig. 260).

CLASS II

Cyanophyceae, Blue-green Algae (')

The Cyanophyceae are simply organised unicellular or filamentous

Thallophytes of a bluish-green colour ; the cells or filaments are
z
338 BOTANY

frequently united into colonies by the gelatinous swelling of the cell

walls. The numerous species, which are distributed over the earth,

FIG. 261. Gloeocapsa polydtrmatica. Fio. 262. A, Oscittaria pHnceps: a, terminal portion of a
A, In process of division ; B, to filament b, portions from the middle of a filament,
;

the left, shortly after division ; properly fixed and stained ; t, cells in division (x 1080).
C, a later stage, (x 540.) B, Oscillaria Froclichii (x 540).

live in water, or form gelatinous or filamentous growths on damp

soil, damp rocks, or the bark of trees.
The protoplast of each cell possesses a peripheral chromatophore of the form of
a hollow cylinder or hollow sphere in addition to ;

chlorophyll this contains a blue green pigment

-

phycocyan from which the name of the class is

derived. The product of assimilation is glycogen.
The centre of the cell is occupied by the colourless
central body, which corresponds to a nucleus and
in vegetative cells is in a condition of continuous
mitotic division. As definite inclusions of the cells
may be mentioned the cyanophycin granules, which
serve as reserve material and are situated in the
neighbourhood of the transverse septa or within
the chromatophore, and the mucilage spheres which
appear in close proximity to the nucleus. Chitin
has been shown to be present in the cell wall.
Reproduction is exclusively vegetative by cell
division. In many forms resting spores are formed
A by the enlargement of single cells, the walls of
Fio. 263. Nostoc LincUi.
which become greatly thickened (Fie. 263). Resting
ment with two heterocysts (h) , .

andalargenumberof spores(<i);
nuclel *** Present ln these S P reS '

B, isolated spore beginning to

J us t as the
Bacteria are designated Fission-
germinate; C, young filament Fungi (Schizomycetes), the Blue-green Algae may
developed from spore. (After jj e termed Fission-Algae
(Schizophyceae), since the
reproduction of both depends on fission. The two
groups would form the class of fission plants Schizophyta. The Bacteria and the
Cyanophyceae have much in common, but the cilia and endospores of the former
 CRYPTOGAMS 339

are unknown in the latter group. According to the observations of PHILIPPS,

which, however, require confirmation, some of the Cyanophyceae which exhibit
movements possess cilia.
The simplest Cyanophyceae consist of spherical cells this is the case with
;

species of Ckroococcus. In Gloeocapsa (Fig. 261), found on damp rocks and Malls,
the cells remain connected together after division into a gelatinous mass, forming
a multicellular colony.
The species of Oscillaria, which occur everywhere in water or on damp soil,
are the simplest of the filamentous forms. The filament, which is usually provided
with a thick sheath, consists of similar flattened cells (Fig. 262). It can separate
into pieces. (hormogonia), which become free owing to the pressure of the sheath,
and grow into new filaments. In other filamentous Cyanophyceae specially modified
cells with their contents degenerated occur in the filament. The significance of
these heterocysts is not yet clear. The species of Nostoc (Fig. 263), whose bead-
like filaments are united by the swell-
ing of the cell-walls into more or less
spherical gelatinous colonies living on
damp soil or in water, afford an example
of this.
ilany Cyanophyceae take part with
the Fungi in the formation of Lichens.
Some species also are endophytic and
inhabit cavities in other plants, e.y.
Anabaena in Azolla, Nostoc in some
Liverworts, in Lemna, and in the roots
of Cycas and Gunnera.

CLASS III

Flagellata (Flagellates)

The Flagellata
are a group of uni-

organisms exhibiting
cellular, aquatic
a wide range of form they combine
;

animal and vegetable characteristics,

and may be regarded as the starting-
point on the one side of unicellular
Fio. 204. Eaylena gracilis. A, Form with green
Thallophytes, on the other of the
Protozoa. chromatophores (ch) n, nucleus v, vacuolenml
; ;

red eye-spot g, flagellum.

; B, Hemisaprophytic
The protoplast exhibits contractile form with small green chromatophores. C,
or amceboid movements, and is limited Colourless saprophytie form occurring in nutrient
solution in absence of light. D, Resting cyst of
by a denser protoplasmic layer and
the form C r, red eye-spot. E, Germination of
not by a definite cell wall. One or ;

the resting cyst of the form A by division into

more cilia (flagella) are present as four daughter cells which later escape. (After
motile organs. The protoplast con- /UMSTEIN A,C, x630; B, X650; D, E, xlOOO.)
;

tains a nucleus, a pulsating vacuole,

and in many species well -formed green, yellow, or brownish -yellow chromato-
phores. Other forms are colourless and live saprophytically ; many are also able
to take up solid particles as food.
340 150TANV

,Mi>t representatives of the group live as naked, free cells ; others form more
or less complicated cell-colonies held together by mucilage, or they possess peculiar
stalked or uustalked firm investments.
Sexual reproduction
is wanting. Multiplication takes plaee by longitudinal
division, and species thick-walled resting spores or cysts are produced.
in many
Euyli'H" Kig. 264) may be taken as an example.
'

The speeies live in water

containing organic substances and often appear in large numbers. By varying
I lie nutritive conditions colourless forms \\itli reduced chnmiatoplimes are
8
obtained ( ).

CLASS IV

!>

Myxomycetes (Slime-Fungi) ( )

The Myxomycetes form an independent group of lower Thallo-

phytes ;
in certain respects they occupy an. intermediate position
between plants and animals, and have in consequence also been termed
Mycetozoa or Fungus-animals. They are represented by numerous
species, and are widely distributed over the whole earth. In their
vegetative condition the Slime-Fungi consist of naked masses of proto-
plasm, the FLASMODJA, containing numerous small nuclei but utterly
devoid of chlorophyll. Glycogen occurs as a reserve substance, while
starch is not found. The plasmodia (p. 54) are found most frequently
in forests, upon soil rich in humus, upon fallen leaves, and in decaying
wood. They creep about on the substrata, changing their form at the
same time, and thrust out processes or pseudopodia, which may in
turn coalesce. At the period of spore-formation the plasmodium
creeps out from the substratum towards the light and air, and, after
coming to rest, is converted into single or numerous and closely
contiguous fructifications, according to the genus. On the periphery
of each fructification an outer envelope or I'KRIDIUM is formed while ;

internally the contents of the fructification separate into spores, each

of which is. provided with a nucleus, and enclosed by an outer wall.
The isolation of the spores may take place on the conclusion of cell
divisions (Trirhia) ; in other cases (e.g. Fuligo) the plasmodium divides
into a number of multinucleate portions while nuclear division is in
progress, and these portions subsequently divide, giving rise to
the spores. The spores thus formed have accordingly an asexual
origin. In many genera, part of the internal protoplasm within the
SPORANGIUM or spore-receptacle is utilised in the formation of a
CAPILLITIUM (Fig. 266 />'), consisting of isolated or reticulately
united threads or tubes. Upon the maturity of the spores, the
peridium of the sporangium becomes ruptured, the capillitium expand.-
(Fig. 265 J5), and the spores are dispersed by the wind, aided by the
hygroscopic movements of the capillitium. In the case of the genus
'iTiifiomi/.ra, the process is somewhat simplified, as the fructification
i

is not
enveloped by a peridium, and the spores are produced at the
SI-XT. I CRYPTOGAMS 341

extremities of short stalks, SKXTAL UKI-KODUCTION is entirely absent

in the Myxomycetes.

A good example of the development of the plasmodia from the spores is afforded
ly Chondrioderma '////;-/////, a Slime-Fungus, common on decaying leaves, dung,
etc., upon which it forms small, round, sessile sporangia. The germination of the
-pores (Fig. 59 a) may be easily observed when cultivated in an infusion of
(.'abhage leaves or other vegetable matter. The spore-wall is ruptured and left
empty by the escaping protoplast. After developing a flagellum or riLU'M as an
organ of motion, the protoplast swims about in the water, being converted into
a sw.viiM-.si'OUK (Fig. 59 e-g), with a cell-nucleus in its anterior or ciliated end,
and a contractile vacuole in the
posterior end of its body. Eventu-
ally the cilium is drawn in, and the
swarm -
spore becomes transformed
into a MYXAMOKiiA ;
these have the
i-apacity of multiplication by division.
In conditions unfavourable for their
development they surround them-
selves with ,i wall, and as MITKIM Y.vrs

pass into a state of rest from which.

under favourable conditions, they
emerge as swarm spores.
-
again
Ultimately a number of the myx-
amoebae approach close together (Fig.
Kio. 265. Ri)' t'ructilications, after listharj:c of tho i

59 I) and coalesce, forming small

spores. -I. Sti'tnonitit fusctt (x 10); An-i/rin 7,',

plasmodia (Fig. 59 m), which in turn <w (x I'-') '', Ci-ibrarin rufc (x 32).
:

fuse with others into larger plasmodia

(Fig. 59 n) no nuclear fusion accompanies this coalescence. Both the amoebae and
;

plasmodia are nourished by the small food particles taken up by the protoplasm,
which also exhibits active, internal, streaming movements. After an interval of
a few days the plasmodium
creeps to the surface of the substratum, and passing
into a resting stage becomes at length converted into a white sporangium with a
double wall, consisting of an outer, calcareous, brittle peridium and an inner
and thinner enveloping pellicle which, in addition to the numerous spores, encloses
also a poorly developed capillitinm.
The development of the other Myxomycetes is -accomplished in a similar
manm-r. Very large plasmodia, often over a foot in breadth, of a bright yellow
colour and creamy consistency, are formed by Fuliyo rarians (Aethnfinnt wjiticum),
"
and as the "flowers of tan are often found in summer on moist tan bark.
If exposed to desiccation, the plasmodia of this Myxomycete pass into a
resting state, and become converted into spherical or strand-like SCLEIKITIA,
from which a plasmodium is again produced on a further supply of water.
Filially, the whole plasmodium becomes transformed into a dry cushion or
cake-shaped fructification of a white, yellowish, or .brown colour. The fructi-
fication, in this instance, is enveloped by an outer calcareous crust or rind, and
is subdivided by numerous internal septa. It encloses numerous dark violet-
coloured spores, and is traversed by a filamentous capillitium, in which are
dispersed irregularly-shaped vesicles containing granules of calcium carbonate.
A fructification of this nature, or so-called fethalium, consists, therefore, of a
number of sporangia combined together, while iu most of the Myxomycetes the
342 BOTANY

and myxamoebae of
sporangia are simple and formed singly. The swarm-spores
the Myxomycetes indicate an origin of the group from organisms resembling
the Flagellata.
The structure and nature of the sporangia afford the most convenient means of
distinguishing theditlVi
-

ent genera. The usually

brown or yellow spor-
angia are spherical, oval,
or cylindrical, stalked
KK;. 267.
(Figs. 265, 267) or not I.eomrpt's fragills.

stalked (Fig. 266). They Groups of sporangia upon a

Moss. (Nat. size.)
usually open by the rup-
ture of the upper portion of the sporangium wall.-,
the lower portion persisting as a cup (Figs. 265
B, 266 A). In Cribraria (Fig. 265 C) the upper
part of the wall of the sporangium, which contains
no capillitium, becomes perforated in a sieve - like,
manner. In Stemonilis (Fig. 265 A) the whole peridium
falls to pieces, and the capillitium is attached "to a

collumella, which forms a continuation of the stalk.

10
Plasmodiophora Brassicae ( ), one of the few-
parasitic Myxomycetes, causes tuberous swellings on
the lateral roots of various species of Brassico. Its
multinucleate myxamoebae occur in numbers in thecells
of the hypertrophied parenchyma of these swellings ;
after the contents of the host-cell have been ex-
hausted they fuse into plasmodia, and these, eventu-
ally dividing into numerous spores, are set free by
the disorganisation of the plant. According to PRQ-
WAZEK the plasmodium first divides into unimicleatc
cellswhich then conjugate in pairs to give rise to
the spores the nuclei before the fusion have under-
;

FIG. 266. Trichia varia. A, gone a reduction division. The spores germinate like
Closed and open sporangia
those of Chondrioderma, and the myxamoebae penetrate
(x 6) B, a fibre of the capil-
;

litium ( x i!40) C, spores (x

the roots of a young Cabbage-plant. The formation
;

240).
of true sporangia, however, does not take place, and
this Slime-Fungus represents a more simply organised
or, in consequence of its parasitic mode of life, a degenerate Myxomycete.

CLASS V
Peridineae 6 -
n
( )

The Peridineae or Dinoflagellata are connected as an independent and further

developed group with certain Flagellata (Cryptomonadinae). They occur as
unicellular, free-swimming organisms in fresh water, but for the most part in the
sea where, together with the Diatomeae, they constitute an important constituent
of the plankton. Their cells are characterised by the possession of two long cilia
or flagella which spring from the middle of the ventral surface in a longitudinal
furrow one of the cilia is directed backwards, the other is thrown into t-urvt- .-tin
;
I
 CRYPTOGAMS 343

lies in a transverse furrow (Fig. 268 A, <//). The protoplast contains a nucleus,
vacuoles of different sorts, and numerous brownish-yellow chromatophores ; the
latter contain a mixture of several special pigments (reddish-brown phycopyrin,

in
schl
FIG. 208. A, Gymnodiniumrhomboides: If, longitudinal groove gf, transverse groove. B, Goniodoina
;

acuminatum the flagella, which arise from the orifice gsp, are not represented schl, d, plates
: ;

of the wall g, girdle.

; C, Peridinivm tabulatum h, wall.
:
(A, B, After SCHUTT C, after ;

KLEBS from OLTMANN'S Algae.)

;

red peridinin, and yellowish-green chlorophyllin). Oil is usually formed as the

product of assimilation. While the Gymnodineae (Fig. 268 A) have either naked
cells or cells limited by a uniformly thickened cellulose wall, the typical Peridineae

Fin. 269. Peridineae from the plankton. A, Ornithocerctts splendidttg ( x 150). B, C, Species
of Ceratium ( x 125.) (After SCHOTT.)

have a wall composed of cellulose formed of polygonal plates these are usually ;

delicately sculptured and perforated with pores. The transverse furrow is formed
by one girdle-shaped plate (Fig. 268 B}.
344 BOTANY

many Peridineae of the plankton the plates bear special wing-like expansions
In
(Fig. 269 A) or the cells have long horn-like processes. These adaptations enable
the organisms to remain floating in the water ( 12 ).
In some Peridineae the chromatophores are only represented l>y colourless
leucoplasts. Such species live either as saprophytes or in the same way as
animals. iliiniiiiMliii! inn hyaliiiuin is a colourless, naked, fresh-water Conn, the

protoplast of which for the purpose of absorbing nourishment loses its cilia
and assumes the form of an amceba in this condition it encloses and digests
;

1:|
small Algae ( ).

Reproduction is by cell-division and also by means of swarm-spores. In the

latter case the protoplast escapes from the membrane and either shortly Conns a
new wall or divides into several swarm-cells (Fig. 268 C}. Thick-walled resting
cells (cysts) also occur. ZEDERBAUKK has recently observed a process of sexual
reproduction (the conjugation of protoplasts escaping from two similar cells to
form a zygote) in Ceratium ( 14 ).

CLASS VI

ls
Conjugatae (" )

This class of green freshwater Algae includes unicellular and

simply filamentous forms, and is clearly distinct from that of the
Chlorophyceae. The Conjugatae
are most nearly related to the
Diatomeae, with which they might
be united in the group of the
since the two classes
sn w essential agreement in their
fZygophyceae,
^B^fc-
yj^p
ir^r*
J&Sb. /^dJEB-hT^.
:{fffP f }
mode of sexual reproduction. This
consists in the conjugation of two
\J u equivalent gametes to form a xygote
or zygospore. Asexual reproduction
,

ating cells each divided into two daughter by means of SWarm-SpOl'eS is Wai iting
cells 2, S, stages of conjugation
; mature j n Jj ot h claSSeS.aild Since the (SimeteS
; ,

zraotes. (After ABCHXR from OLTMANN'S : , .,, .,. ., .

, -,

jigae.)
also are unprovided with cilia these
forms are also classed as Acontae.
The cells of the Conjugatae which increase in number by cell-division
are uninuclear, and differ from those of the Diatoms in having a cell-
wall which is not silicified and in the
presence of large green chloro-
plasts of complicated structure.

1. The Mesotaeniaceae, the

simplest of the unicellular Conjugatae, include only
a few genera. They are
distinguished from the following order by the cell-wall of
the shortly cylindrical cells not
being formed of two halves. The mode of eon
jugation presents some differences. In :>',///<. /^/w (Fig. 270) the protoplasts of
the conjugating cells first divide and the cells unite in to form the
daughter pairs
 CRYPTOGAMS :J4f.

zygotes; in Mcsotacii-inm. and 'i/lindrocystis the protoplasts conjugate without a

'

preliminary cell-division. .Similar differences are shown by the Diatomeae. The

zygote on germination divides into four new individuals.
'1. The Desmidiaceae are unicellular or their cells are united in rows they are ;

of great beauty and, like the Diatoms, exhibit a great variety of form. Their
cells are composed of two

symmetrical halves, separated,

as a rule, from each other by
a constriction. Each half
contains a
large, radiate

chromatophore, or a number
of plate - like chromatophores
united into one. Within the
chromatophores are disposed
several pyrenoids, while the
nucleus lies in the centre of
the cell in the constriction.
The cells themselves display
a great diversity of form and
external configuration, being
sometimes rounded (e.g. COK-
marium, Fig. 271 A, B) some-
times stellate (Micrasterias,
Fig. 271 ]'>). The cell walls,
which, as in the Diatoms, con-
sist of separate halves, are
frequently beset with wart- or
horn-like protuberances and
often provided with pores. In
some genera there is no con-
strictionbetween the two
halves of the cell. This is the
case, for instance, in the cres-
cent-shaped Closterium monili-
/'/ /?i( Fig. 272 F), whose two

chromatophores consist of six

J)
elongated plates, united in
the long axis of the plant, Kiii. _'. I. .1. I'IIXHIII fin HI i-oelatuni , dividing. B, C,
while in each end of the cell liotri/tis. Cj, two cells at right angles prepariii for con-

there is a small vacuole con- jugation the lower cell shows the conjugation canal ('.,,

minute of gametes fused into the young zygote ; Cy, mature />;
taining crystals //, .U ii'i'usii riii-- t'l-ii, iiirliteiisix. (After RAI.KS C] :

gypsum in constant motion. alter DK JURY.)

Many Desmids are character-
ised by heliotactic movements they protrude fine mucilaginous threads through
;

the cell walls, by means of which they can push themselves along, and take up a
position in a line with the direction of the incident rays of light.
Multiplication is effected by cell-division. This is accomplished by the forma-
tion of a partition wall across the middle of the cell after the nuclear division is

completed. Each daughter cell eventually attains the size and form of the mother
n 11.
by the outgrowth of a new half on the side towards the new division wall
(Fig. 271 A}.
346 BOTANY

The conjugation of the protoplasts takes place, in the case of the Desini-
diaceae, outside their cell-walls. Two cells approach each other, and surround
themselves with a mucilaginous envelope. Their cell-walls rupture at the con-
striction, and parting in half allow the protoplasts to escape ; these then unite
to form a zygospore. The zygospores frequently present a very characteristic
appearance, as their walls are often beset
with spines (Fig. 271 C). The four empty
cell halves may be seen close to the
spore.
In some
Desmidiaceae the conjugating
cellsundergo a preliminary division, the
daughter cells uniting in pairs.
The two sexual nuclei in the zygote do
not fuse until germination of the latter is
about to commence. The resulting nucleus
then undergoes division into four nuclei,
^
* two large and two small. Only two cells
are formed from the zygote each of which
has thus two nuclei of different sizes ;

the smaller nuclei disappear. The pro-

duction of two cells on germination thus
appears to be derived from the division
into four cells seen in Mcsotaenium, and to
stand to the latter as a reduced form.
3. Zygnemaceae. In this family, all
of which are filamentous in character, the
genus Spirogyra, with its numerous species,
isthe best known. It is commonly found
in standing water, forming unattached
masses of tangled green filaments. The
filaments exhibit no distinction of base
and apex, and are composed of simple
rows of cells, which vary in length in differ-
ent species. Growth results from the
division and elongation of the cells in
one direction only. Each cell has a large
nucleus and one or several spiral green
band-like chromatophores (Fig. 273 C),
Fio. 272.-Oosfcrti<i. A, Zygote before ger- The cells of Zygnema contain two star-
mination showing the two nuclei not yet
gh d chromato r hores .
united ; B, germinating zygote with the *L .

nuclei united ; C, division into two cells CONJUGATION, in the case of Spirogyra,
each containing one larger and one smaller is preceded by the development of con-
nucleus D, further state of germination ;
;
verging lateral processes from the cells
E, young plants escaping from the cell- of adj acent filaments. When two pro-
membrane; F, Closterium moniliforme ,

mature plant. (A-E after KLEBAHX.)

cesses from opposite cells meet (Fig.
273 A), their walls become absorbed at the
point of contact, and the whole protoplasmic contents of one cell, after contracting
from the cell wall, passes through the canal which is thus formed into the opposite
cell. The protoplasm and nuclei of the conjugating protoplasts then fuse
together while the chloroplasts do not unite, but those of the entering protoplast
disorganise. The resulting cell forms the zygospore invested with a thick wall,
and filled with fatty substances and reddish-brown mucilage-spheres. This form
SECT. I CRYPTOGAMS 347

of conjugation, which is the one exhibited by most species, is described as scalari-

form (Fig. 273 A), as distinct from the lateral conjugation of some species, in
which two adjacent cells of the same filament conjugate by the development of
coalescing processes, which are formed near their transverse wall (Fig. 273 ). In
some genera the zygote is formed midway in the conjugation tube.

Fio. 273. A, Conjugation of Spirogyra qulnina (y. 240). B, Spirogyra longata(x 150); *,
C, Cell ofSpirogyra jugalis k, nucleus
; ; ch, chromatophores ; p, pyrenoid. (x 250.)

On germination the zygote gives rise to only one plant its contents grow ;

out as a cylindrical cell which on division forms a filament. According to

CHIMIELEWSKY the nucleus of the zygote of Spirogyra first divides amitotically into
four, two of which disintegrate while the remaining two again fuse to form the
16
single nucleus of the young plant ( ).

CLASS VII

1T
Diatomeae (Diatoms) (" )

The Diatomeae (Bacillariaceae) constitute a very large class of

unicellular Algae. They occur usually associated together in large
numbers, in both fresh and salt water, and also on damp soil.
The cells are either solitary or form colonies ; they are free-swim-
ming, or are attached by means of gelatinous stalks, excreted by
the cells themselves (Fig. 274). Sometimes the cells remain con-
nected and form bands or zigzag chains, or, on the other hand, they
are attached and enclosed in gelatinous tubes, while in the case of
34S BOTANY

the marine genus Schizonema they lie embedded in large numbers

in a gelatinous branching thallus, often over 1 dcm. in size. The
cells also display a great diversity of shape while generally bilater- ;

all\r symmetrical, they may be circular or elliptical, rod- or weil^e

shaped, curved or straight. The

structure of their cell walls is
especially characteristic ; it is com-
posed of two halves or VAI.YKS,
one of which overlaps the oilier
like the lid of a box (Fig. 3. />').

The present two alto-

cells thus
gether different views, according to

I-' in. -JT'i. Formation ( iiiixosi>uivs in

i-ii-'iiliiln. J. Cell seen from tlie valve side.

/;. Two cells lyinj: alongside one another : their
((intents have divided btto two danghtor eells,
.ach of which ]>ossesses two nuclei, i
. /'. C,,n-

.iu.^atinn in pain of the daughter calto to inrm

tlie auxosiHires, wliich at lirst contain four

Kiu. -274. I. icmophoru flubelln In. nuclei. E, The two t'ull-j;iown .-uixosiMiies ; tin-

Colony of Diatoms with liranched two larger nuclei in each have fuseil into one,
tin- two .smaller ones have disintegrated. (After
gelatinous stalks. (After SMITH,
i'roni GOEBEL'S OrganofrapMe.) N U:STKX, x 500.)

the position in which they are observed, whether from the .IKDLK
(Fig. 3 B] or VALVE SIDE (Fig.
3 A}. The two valves are so strongly
impregnated with silica, that, even when subjected to intense heat,
the original form and
they remain as a siliceous skeleton, retaining
of the cell walls. The walls of the cells, particularly cm
markings
the valve are often ornamented with numerous fine, transverse
side,
 CRYPTOGAMS 349

markings or ribs, and also with small protuberances and pits, or

they are perforated by open pores which serve to give exit to the
gelatinous secretion and possibly play a part in the movements
sometimes exhibited by the cells.

According to the structure of the cell walls two main groups of Diatoms are
distinguished, the Centricae (Fig. 277) with usually circular valves and radial or
concentric sculpturing, and the Pennatac the valves of which are usually elongated
and the sculpturing pinnate. In many of the Pennatae (Fig. 3) a longitudinal
line corresponding to an opening in the cell walls, and exhibiting swollen nodules
at both extremities and in the middle, is distinguishable in the surface of the
valves. Forms provided
with such a median suture "
or i: A THE
XS^ZZTx.
are characterised
^^
/"'"~" ^"v
7
.^TIT^. l/b /^fes. A '-//*.

by peculiar backward-creep-
ing movements, resulting
from the extrusion of pro-
toplasmic protrusions from
the spirally arranged longi-
tudinal slits which are pre-

sent in the raphe (cf. p. 266).

The
has always
cell
a central nucleus and
one (Fig. 3) or two to
four (Fig. 275) large or
numerous smaller chro-
matophores (Fig. 277)
embedded in its parietal

protoplasm. These
chromatophores are flat,

frequently lobed, and

of a brOWnish-yelloW I-'ic. STO. Coccone'is yliuxutulv. 1, Vegetative cell pair of ; ..'.

ce " s M ore conjugation J,

' '' cells in process of conjugation.
>

colour Globules Of a
'
; t
,
,

r.art;e nucleus;
. ijl,-,
small nucleus; /,/,-,gelatinous sub- 11,

tatty Oil are alSO 111- stance. (Alter KAHSTKN. from OI.TMASNS' .!?</" .)
eluded in the cell con-
tents, and take the place of starch as an assimilation product.
The Diatomeae multiply vegetatively by longitudinal division which
always takes place in one direction. In this process the two valves are
firstpushed apart from one another by the increasing protoplasmic
contents of the mother cell, which then divides longitudinally in
such a direction that each of the two new cells retains one valve of the
mother cell. After the division of the protoplasm of the mother cell
is accomplished, each daughter cell forms, on its naked side, a new
valve fitting into the old one. The two valves of a cell are therefore
of different ages. In consequence of this peculiar manner of division,
as the walls of the cells are silicified and incapable of distension, the
daughter cells become successively smaller and smaller, until finally,
after becoming reduced to a definite minimum size, they undergo
 r
3. ,0 BOTANY

transformation into AUXOSPORES. The auxospores are usually two

or three times larger than the cells from which they arise, and by
their further development they re-establish the original size of the
cells.

The formation of the auxospores, which correspond to the zygospores of the

Conjugatae, is accomplished in various ways.
The process in Navicula (Fig. 275),
Pleurosigma, etc. may be compared with the zygospore formation
of Spirotacnia ;
,

two cells lay themselves side by side, the protoplast of each undergoes division
into two, and the resulting cells emerge from the valves of the cell wall and
conjugate in pairs. By division of the nucleus each gamete was provided with a
larger and a smaller nucleus in the auxospores the two large nuclei fuse, while
;

the two small nuclei disintegrate.

In Surirella and Cocconds (Fig. 276) the conjugating cells do not, undergo
division, but unite directly with one another.
The nuclei, however, divide ; in
the former two nuclear divisions occur in each of the conjugating cells,
genus

Fio." 277. Planktoniella sol. 'A disc-shaped Fio. 278. Bacti *i-ifti -i varians. From the
Diatom from the Plankton. A parachute-like, Plankton. A chain of cells which Ix-ar
hollow [membranous wing springs from the bristles as a floating apparatus. (After
girdle. (After SCHUTT. x 190.) Scathr. x 200.)

resulting in one large and three small nuclei in the latter genus there is only
;

a single nuclear division giving one large and one small nucleus. The large
nuclei fuse, the small ones degenerate.
In Achnanthes thecontents of a single cell divides into two daughter cells,
cell
which escape, and then
fuse together to form the auxospore.
In many genera sexuality is not obvious in the process of forming the auxospores,
which arise asexually from cells that do not conjugate. This is the case in
Synedra and Rhabdonema, in which a cell divides and the two daughter cells
emerge and develop into auxospores. In Melosira there is no longer any cell
division, the protoplast enlarges, bursts the cell wall, and becomes directly trans-
formed into an auxospore. The original preliminary divisions are, however, still
frequently indicated by nuclear divisions, in Melosira only by the appearance of
two nucleoli in the nucleus of the auxospore.
Some plankton Diatoms exhibit a breaking up of the protoplast by repeated
 CRYPTOGAMS 351

cell Division into numerous small, round, naked daughter cells. According to
KARSTEN'S observations in Corethrbn, these cells escape, conjugate in pairs, the
resulting zygotes increase in size, and ou germination give rise to two daughter
cells. Each of the latter has at first two nuclei, one of which later disappears,
and the cell grows into a mature individual of Corethron. These processes call to
18
mind the behaviour of Closterium among the Desmidiaceae ( ).

Countless numbers of Diatoms live in the ocean, and they constitute also a
12
proportionately large part of the PLANKTON ( ), that is, the free-swimming organic
world on the surface of the sea. The plankton Diatoms are mostly centric forms,
have no middle suture or raphe on the surface of their valves, and are especially
adapted to swimming or floating. To this end they are often provided with horn-
like protuberances or membranous wings, which greatly enhance their buoyancy

(Figs. 277, 278).

Many Diatoms occur in places where decomposing substances are present in
abundance. Such species can assume a saprophytic mode of life, their chromato-
phores becoming colourless and reduced in size. It has been shown that some
colourless species of Nitzschia which occur in the sea are exclusively dependent
on organic substances for food, the reduction of their chromatophores being
19
complete ( j.
Diatoms occur also as fossils. Their silicified valves form a large part of the
deposits of SILICEOUS EARTH (Kieselguhr, mountain meal, etc.,) and in this form
they are utilised in the manufacture of dynamite.
On account of the extreme fineness of the markings of their valves, it is
customary to employ certain species of Diatoms as test objects for trying the
lenses of microscopes. Pleurosigma angulatum is commonly used for this purpose.

CLASS VIII

Heterocontae ()
In the Heterocontae a number of genera of green Algae are included which were
formerly placed in the Chlorophyceae but are now separated as an independent class.
At the base of the class may be placed
such true Flagellata as Chloramoeba hetero-
morpha (Fig. 279), which lives in fresh
water. The naked, amoeboid cell contains a
nucleus and 2-6 yellowish-green chloroplasts ;

at the anterior end, just outside a vacuole,

two cilia are situated which are of very un-

equal length. It is this last feature which

gives the name to the group. Chloramoeba

is one of those low green organisms which

become colourless and lose the power of

independent nutrition when cultivated in a
nutritive solution in the dark. Thick-walled Fio. 279. Chloramoeba heteromorpha ?,:

resting cells also occur. vacuole k, nucleus ; 3, a resting <''!!-

;

Among the higher Heterocontae char- (After BOHLIN, from OLTMANNS' Algae.)
acterised by the presence of pectin in the
cell wall the genus Conferva which is
widely spread in fresh water must be
mentioned (Fig. 280). The plant consists of simple unbranched filaments the
 BOTANY

cells of which have peculiarly constructed walls; the wall run.-iM- of two parts
separated by an oblique annular split at the middle p.irt of the cell. On cell
division a new portion, H-sliaped in longitudinal section, is intercalated. Tin
characteristics of the group are seen in the yellowish-green, oil-forming chloro-

plasts,and the formation of zoospores with cilia of unequal lengths in many

Confervae the zoospores lum- only a single cilium). In addition to zoospores thick -

walled aplanospores arise by

the separation by the cells
"I' the filament.
With some reservation
OI.TMANNS also places here
ll<il i-
ijil lii-i>i ij
rii /mint ti, IL (Fig.
281). which was formerly in-
cluded in the Siphoneac.
Tliis Alga grows on damp

Kii;. -Jsn. .-fii,ij,',-i-n lii-nilii/i-hif. 1, Filament : -'. '. formation

of transverse wall (</) in cell division; i>, formation <>t"

aplann.sixirfs by lireakinx down of tin- fllainent ; 1<>. Pitt -Si. I. /'. M

Xoos] HUTS with cilia of unequal length. (After CAY /at H ni. .'., The \v li<ili> plant
:

(/. .'), lion LIN (.'. :.'). l.i TIIF.R (/"). From OI,IMAS\>' /.', S\varni-s|Kni-. (.1 x 28:
Algue.) /: .MO.)

clayey soil, where it forms groups of green, balloon-shaped vesicles about '2 nun.
in breadth. These are attached to the soil by branched colourless rhixoids.
The whole plant corresponds to a single multinucleate cell : its protoplasm
contains numerous green chloroplasts. zoospores, produced in large number>
The
by the division up of the contents, escape by an opening at the summit, ivu'h
has a single cilium and contains two chloroplasts. After swarming the S|H,I,
surrounds itself with a wall and grows into one of the balloon-shaped plants (*).
In some genera, gametes resembling the zoosp<nvs but conjugating in pair*
have been observed.
SECT. I CRYPTOGAMS 353

CLASS IX
-"' 21
Chlorophyceae (- )

When the green Conjugatae and Heterocontae are separated there

remains the natural group of the Chlorophyceae, including several
series of genera. The majority of these Algae live in fresh water
or in damp situations, but a large number are found in the sea.
Their characteristic chloroplasts are of a pure green colour, frequently
contain pyrenoids, and nearly always form starch. The asexual
swarm-spores are pear-shaped, and in typical forms possess two or
four cilia of equal length (on this account the group is sometimes
termed Isocontae) and a curved or bowl-shaped chloroplast. In some
genera the swarm-spores are replaced by non-motile aplanospores, and
in certain of the more advanced genera (Oedogonium, F'aucheria) the

swarm-spores are of more complicated structure, but can be derived

from the typical simple form.
Of the five orders included in the Chlorophyceae the Volvocales
stands nearest to the Flagellata and, as is also the case with the
Protococcales, include unicellular and colonial forms. The Ulotrichales
and Siphonocladiales are filamentous ; in the former the filaments are
composed of uninucleate, in the latter of large multinucleate cells.
The filaments are simple in the lower forms, but branched in the
more advanced ones. The thallus of the Siphonales is formed of a
single multinucleate cell.
In all the orders sexual reproduction is
usually effected by the
conjugation of gametes which re-
semble the zoospores. In all the
groups, except the Protococcales,
isogamy is replaced by oogamy
in the higher forms.

Order 1. Volvocales. Typical re-

presentatives of this order are charac-

terised by the cilia being retained by
their cells in the vegetative stage the ;

plants are therefore motile. Each cell

has a nucleus and a chloroplast. The
Volvocales thus resemble the Flagel-
lata, some forms of which, such as Poly-
blepharis, might, on the ground of their
cell structure, be placed in the former FIG. 282. 1. Chlamydomonas angulosa. (After
DlLL -> ' cilia v vacuole *. nucleus chr,
group, but .differ by the absence of a
= > ;

,,,, i^-
cell wall and of sexual reproduction.
chromatophores py, pyrenoid. 2. Polytoma
;

uvdla (after DANGEARD) a> eye-spot. (From.

Clilamydomonas (Fig. 282) and OLTMANNS' Algae.)

Hacmatococcus (Fig. 283) are widely dis-
ffl
tributed forms consisting of free-swimming cells ( ). In the former the cell
2A
354 BOTANY PART II

membrane is closely applied to the protoplast, at the anterior end of which two cilia

Fio. 283. B, Haeinatococcus plurialis (x 360) ; A, swarming cell ; ]',, formation of swarm-
^4,

spores. C-G, Baematococcus Biitschlii ; C, formation of gametes ( x 400) ; D, gamete :/..

conjugation of two gametes t', <!, xygotes(x 800). (<'>!. alt*

; iHi u. MMAVN.)

Fio. 284. Volvox globator. A, Colony showing various stages of development of ova. and
spermatozoids (x 165). B, Bundle of spennatozoicls formed l>y division from a single
eell(x 530). C, Spermatoxoids ( x 530). D, Kgg-cell surrounded by spermatozoids in
the mucilaginous membrane (x 26o). (After F. COHN.)

and a red eye-spot are situated ;

in the latter genus the membrane is separated from
 CRYPTOGAMS 355

the protoplast by a gelatinous layer except at the anterior end. Haematococcus

pluvialis occurs commonly in puddles of rain-water, and, like Chlamydomonas
nivalis, which gives rise to "red snow" and occurs on snow in the Alps, etc.,
is characterised by the presence of a red pigment (haematochrome) in the cells.

Reproduction both asexual, by swarm-spores, 2-8-16 of which are formed in a

is

mother cell and

are set free by rupture of the membrane, and sexual the sexual ;

reproduction is by conjugation of similar, small, biciliate gametes formed in large

numbers (to 64) in a mother cell, and uniting in pairs by their anterior ends to
form a zygote. In Chlamydomonas coccifera, according to GOROSCHANKIN ( 23 ),
there is in contrast to the other species a marked differentiation in the sexual
cells. Single cells become transformed into large, non-ciliated, female gametes or
egg-cells others divide and each gives rise to 16 small, biciliate male gametes.
;

The transition to oogamy thus occurs in this group even among the isolated
unicellular forms.
Polytoma uvella, which resembles Chlamydomonas in structure, is a colourless
and saprophytic form (Fig. 282, 2).
The biciliate cells of Pundorina, Eudorina, Volvox, etc., are united in colonies
or coenobia. In Volvox ( 24 ) (Fig. 284), which may be regarded as the highest
form in the order, the free-swimming colonies have the shape of a hollow sphere.
The component protoplasts are connected by fine processes, so that the organism
must be regarded as constituting a single individual. The sexual cells are
differentiated into ova and spermatozoids. The egg-cells arise by the enlarge-
ment of single cells of the colony; they are large, green, non- motile cells
surrounded by a mucilaginous wall. The small spermatozoids are elongated
bodies of a bright yellow colour, provided with two cilia at the colourless
anterior end they arise by the division of a cell of the colony into numerous
;

daughter cells. After fusing with a spermatozoid within the cavity of the
colony the egg-cell is transformed into the thick- walled, resting oospore. The
vegetative reproduction of Volvox takes place by the division of single cells of
the colony to form a new daughter colony this corresponds to the formation of
;

swarm-spores in other genera. Eudorina is also oogamous.

Order 2. Protococcales. These are unicellular green Algre, or their cells are
united in colonies of various form the vegetative cells have no cilia, and the cell
;

or colony is consequently non-motile.

Usually each cell contains a nucleus
and only one chloroplast. Reproduc-
tion is by means of zoospores, in place
of which in many genera non-ciliated
aplanospores are found. Sexual repro-
duction, when present, takes the form of
conjugation of similar gametes.
The simplest forms belong to the
genera Chlorococcum (including Cysto-
25 26
coccus) and Chlorella C
'

). The cells FIG. 2S5. Chlorococcum (Chlorosphaera) limicola. 1.

of the former are spherical, and occur Vegetative cell and cell divided into 8 zoospores ;

in fresh water and also on 2, free zoospores 3, zoospores after they have
damp sub-
;

formed cell-walls. (After BEYERINCK, from

strata they frequently take part in
;
OLTMANNS' Algae.)
the composition of Lichen thalli.
Asexual reproduction is by the production from a cell of a number of biciliate
zoospores (Fig. 285) ; under certain conditions these are replaced by aplanospores
without cilia. Chlorella milgaris (Fig. 286) is a wide -
spread Alga, the small
2 A 1
356 BOTANY

cells ofwhich often live symbiotically in the protoplasts of lower animals (Infusoriae,
Hydra, Spongilla, Planariae) it is multiplied only by division of the cells into
;

2, 4, or 8 aplanospores which surround themselves with a wall and grow to the

full size.
The simplest type of cell-colony,
consisting of four cells, is found in the
27
genus Scenedesmus C ), which is

widely spread in fresh water,and

connects on to Chlorclla. The com-
monest form, Sc. acutus, has spindle-
shaped cells, while the colonies of Sc.
caudatus are distinguished by four long
horn-like prolongations of the cell wall
(Fig. 287). In reproduction each cell
divides in the direction of its length
FIG. 286.-CorHa mdgafis. Cell division
1, ; 5, 3, into four daughter -cells, which, on
into four aplanospores U, 5, division into .eight
{ f h u f
aplanospores. (After GRINTZESCO.)
colony. new
More complicated cell
colonies are met with in Pediastrum ( M ) (Fig. 288), in which each cell-family
forms a free-swimming plate, composed internally of polygonal cells, while on the
margin it more or less acutely crenated. The formation of asexual
consists of cells
swarm-spores Pediastrum by the divjsion of the contents of a cell into
is effected in
a number (in the case of the species illustrated, P. granulatum', into 16) of naked
swarm spores, each with two cilia. The
-

swarm - spores, on escaping through the A.

ruptured cell wall (Fig. A, b), are enclosed
in a common envelope. After first mov-
ing vigorously about within this envelope,
they eventually collect together and form

38. Pcdiastrum granulatum. A, An old

cell-family a, cells containing sporrs
: ; i>.

spores in process of extrusion (the other

cells have already discharged their spon-s) :

Flo. 287. A, Scenedesmus acutus. Jl, tin- .-.ame, B, cell-family shortly after extrusion of the
undergoing division; C, Scenedesmus spores C, cell-family 4J hours later. (After
;

(After SENN, x 1000.) AL. BRAI:N, x 300.)

a new cell-family. Pediastrum also possesses a sexual mode of reproduction.

The gametes are all of equal size, and except that they are smaller and are pro-
duced in greater numbers, are similar to the swarm -spores. They move freely
about in the water, and in conjugating fuse in pairs to form zygotes. The further
development of the zygotes into cell-families is not yet fully known.
The life-history of the Water-net (Hydrodictyon utriculatum ( 29 ) is essentially l-
 CRYPTOGAMS 357

similar. It is one of the most beautiful of the free-floating, fresh-water Algae, the
hollow cylindrical colonies being formed of elongated cells united together to form
a many-meshed net.
Order 3. The Ulotrichales exhibit, as compared with the uni-
Ulotrichales.
cellular green Algae, an advance in the external segmentation of the thallus. It
is always multicellular, and, in most of the genera, consists of simple or branched

filaments. The filaments are either attached by a colourless basal cell to the
substratum (Fig. 290 A) or float free. The thallus of the marine genus Ulva
(Ulxa lactuca, SEA LETTUCE) has the form of a large, leaf-like cell surface, and is
two layers of cells thick (Fig. 5, p. 13). In Entcromorpha (Fig. 289) the thallus
is ribbon-shaped, either cylindrical or flattened when young it is two-layered, but
;

later it becomes hollow, the wall thus consisting of one layer of cells. Although
the majority of the Ulotrichales
live in fresh or salt water, a few
aerial forms (Chroolepideae) grow on

stones, trunks of trees, and, in the

tropics, on leaves. To this family

belongs Trentepohlia (or Chroolepus)

Jolithus, often found growing on
stones in mountainous regions.
The cell filaments of this species ap-

pear red on account of the hsemato-

chrome they contain, and possess
a violet-like odour.
The asexual reproduction is
accomplished by the formation of
ciliated swarm - spores. Sexual re-
production is effected either by the
fusion of planogametes, or the sexual
cells are differentiated as non-motile
egg-cells and motile spermatozoids.
Ulothrix zonata x) 290
( (Fig.
A) is one of the commonest fila-

mentous Algae. The filaments of Ulo- FIG. 289. Enteromorpha compressa. (i nat. size.)'.

thrix exhibit no pronounced apical

growth; they are unbranched, attached by a rhizoid cell, and consist of single rows
The asexual reproduction
of short cells ; each cell contains a band-shaped chloroplast.
is effected by means of swarm-spores, which have four cilia (C), and are formed by

division in any cell of the filament. The swarm-spores escape through a lateral
opening (B) formed by absorption of the cell wall, and, after swarming, give rise to
new filaments. The sexual swarm-cells, or planogametes, are formed in a similar
manner by the division of the cells, but in much greater numbers. They are
also smaller, and possess only two cilia (E). In other respects they resemble the
swarm- spores, and possess a red eye-spot and one chromatophore. By the con-
jugation of the planogametes in pairs, zygotes (F-H) are produced, which, after
drawing in their cilia, round themselves off and become invested with a cell wall.
After a shorter or longer period of rest the zygotes are converted into unicellular
germ plants (/), and give rise to several swarm-spores (K), which in turn grow
out into new filaments. Under some conditions the planogametes can give rise
to new plants parthenogenetically without conjugating'. Further, the filaments
can, in addition to the swarm-spores with four cilia described above, produce

2 A 2
358 BOTANY

others of smaller size (micro - zoospores) which resemble the gametes. These
possess four or two cilia, and as a rule die if the temperature of the medium is
above 10 ;
below this temperature they come to rest after a few days and proceed
to germinate slowly. This Alga is thus of interest from the incomplete sexual
differentiation exhibited by its gametes.
31
The genera Oedogonium ( )
and Bulbuchaetc may be quoted as examples <>t

oogamous Ulotrichales. While tlie

thallus of the latter is branched,

the numerous species of Oedoijmi in m
consist of unbranched filaments,
each cell of \vhirh possesses one nu-
cleusand a single parietal chromato-
phore composed of numerous united
bands. The asexual swarm-spores of
Oedogonium are unusually large, and
have a circlet of cilia around their
kinoplasmic, colourless, anterior ex-
tremity (Fig. 291 ). In this case
the swarm-spores are formed singly,
from the whole contents of any single
cell ofthe filament (A), and escape
by the rupture of the cell wall.
After becoming attached by the
colourless end they germinate, giv-

ing rise to a new filament. For the

purpose of sexual reproduction, on
the other hand, special cells become
swollen and differentiated into
barrel shaped oogonia. A single
-

large egg -cell with a colourless

receptive spot is formed in each
oogonium by the contraction of its

protoplasm, while the wall of the

oogonium becomes perforated by an
opening at a point opposite the
receptive spot of the egg. At the
same time, other, generally shorter,
FIG. 290. Ulothrix zonata. A, Young filament with ,, ., ,.
Cells ot the Same Or another fila '
rtmoid cellr(x 300); B, portion of i.a,M,nt with
escaping swarm-spores C, single swarm-spore D,
; ;
ment become converted into an-
fonnation and escape of gametes E, gametes F, G,
; ; theridia. Each antheriditim usually
conjugation of two gametes H, zygote ; J, zygote
;
gives rise to two spermatozoids
after period of rest; A", zygote after division into
The
sperniatozoids are smaller than
swarm-spores. (After DODEL-PORT. B-K x 482.)
the asexual swarm-spores, but nave
a similar circlet of cilia. They penetrate the opening in the oogonium and fuse
with the egg cell, which then becomes transformed into a large, firm-walled oospore.
On the germination of the oospore its contents become divided into four swarm-
which gives rise to a new cell filament. In the adjoining figure (Fig.
spores, each of
292) a germinating oospore of Bulbochaete with four swarm-spores is represented.
In some species of Oedogonium the process of sexual reproduction is more com-
plicated, and the spermatozoids are produced in so-called IHVAKK MAI.KS. These
are short filaments (Fig. 291 C, a) consisting of but few cells, and are developed
.SKOT. 1 CRYPTOGAMS 359

from asexual swarm -spores (.VNDUosj'oiiE.s) which, after swarming, attach them-
selves to the female lilaments, or even to the oogonia. In the upper cells of the
dwarf male filaments thus derived from the androspores, spermato/oids are pro-
duced which are set free by the opening of a cap-like lid (Fig. 291 l>, a).
The genus Coleochacle (*) is also oogamous. The long colourless neck of the
Mask-shaped oogonium opens at the tip to allow of the entrance of the spermatozoid.
The spherical oospore increases in size and becomes surrounded by a single layer
<>f
pseudo-parenchymatotis tissue derived from filaments that spring from the
stalk cell of the oogonium and neighbouring cells. In this way a fruit-like body
is formed. On germination the oospore first divides into 16-32 wedge-shaped
cells, then breaks up and liberates a swarm-spore from each cell. While in regard

BH
1

/;,

I'H.. $01. A, I!, ii' iliHioiiiinu :

A, escaping swiinn -
spores : /;. formation of tour swarni-
tree suMi-iii-spore. <', 1), Onlixjniinnn i-ilintinn : '
',
Ix-fore fer- s]H>res in tin- x'Tiiiiniitinx
tilisation ; 7), in process nl' fertilisation : .
ooj;onia :
n, <!>.v;irf oospore. (After I'KIM;-.-
males : N. s]iiT!M;itn/oiil. (After I'm xcsHKlM, X 350.) II KIM, X '-'"lO.)

to these processes (Jvlcochaete exhibits the highest grade attained by the Green
Algae, cannot be regarded as the point of origin of the Bryophyta.
it

Order 4. Siphonocladiales. The Algae of this order are filamentous and

usually branched they are distinguished from the Ulotriehales by their large
;

multinucleate cells, the chloroplasts of which are either solitary, large and reticn-
lately formed, or appear as numerous small discs.-
The genus I'liiiLn/ilnn-n. numerous species of which occur in the sea and in fresh

water, one of the most important representatives of the order.

is Cl. glomerate

(Fig. 29-'5) is one of the commonest Algae in streams, often attaining the length of
a foot. It is attached by rhizoid-like cells, and consists of branched filaments with

typical apical growth which some other representatives of the order do not show.
The structure of the cells is represented in Fig. 61. Branching takes place from
the upper ends of the cells by the formation of a protrusion which is cut off as the
first cell of the branch. Asexual reproduction is by means of biciliate zoospores
(Fig. 293), which arise in numbers from the upper cells of the filaments, and cse.-ipe
from these sporangia by a lateral opening in the wall. The sexual reproduction is
isogamous as in ( 'fothrix.
Only in the genus Sphneroplca has the sexual reproduction become oogamous.
S. annulinu consists of simple filaments and occurs in fresh water.

Many Conns occur in the sea (e.<j. Sipkonockuhu), and some have a highly com-
plicated thallus. which is always, however, formed of branched filaments by ;

2 A 3
360 BOTANY

calcareous incrustation some forms conic to resemble coral. A'-ihil.tilnrin ni<ii;t< ,

-

ranca (**) example of such calcareous Algae. Tin- thin

(Fig. 294) will serve as an.
stalk of the thallus is attached by means of rhizoids, while the umbrella-like disc
consists of closely united tubular outgrowths, each of which is to be regarded as
a gametangiom. The contents of the latter do not fonn the biciliate gametes
directly, but lirst divide into a large
number of firm-walled cysts. The-,
remain in the resting condition
throughout the winter, and then
give rise to numerous gametes which
conjugate in pairs. The zygotes
germinate promptly and grow into
new plant-.
Order 5. Siphonales. The Siph-
oneae are distinguished from the pre-
ceding groups of Algae by the struc-
ture of their thallus. which, although
more or less profusely branched, is

K 11 :._". '3. Portion of Cladopho, -n ijlo,i>enit. (x48.) l'i<:. 294. Acetiiliii/nriii med
To the ri^lit a swarm-spore, (x 1000.) (\at. size.)

not atfirst divided by transverse septa. The cell-wall thus encloses a continuous
protoplasmic body in which numerous nuclei and small green chloroplasts are
embedded. The same type of thallus is also met with in the Phycomycetes or
Algal Fungi.
The majority of the Siphoneae inhabit the sea, and on account of the com-
plicated segmentation of their thallus, afford one of the most interesting types
of algal development. The genus Caulerpa ( :M ), represented by many species
inhabiting the warmer water of the ocean, has a creeping main axis. Increasing
in length by apical growth, the stem-like portion of the thallus gives off from its
under surface profusely branched colourless rhizoids, while, from its upper side,
 CRYPTOGAMS 361

it produces green thalloid segments which vary in shape in the different species.
In Caulerpa prolifera (Fig. 295) these outgrowths are leaf-like and are frequently
proliferous. In other species they are pinnately lobed or branched. The whole
thallus, however branched and segmented it may be, encloses but one cell -cavity,
which is, however, often traversed by a network of cross-supports or trabeculaj.
Starch-forming leucoplasts are present in the colourless parts of the thallus.
The genus Bryopsis, on the other hand, has a delicate, pinnately branched
thallus. The thallus of Halimeda, the species of which occur in the wanner seas,
is composed of flattened
segments, and resembles an Opuntia on a small scale. By
incrustation with lime it at-
tains a hard, coral-like texture.
The segments are formed of
branched tubular filaments.
In Jji'i/opsisthe conjugating
gametes are differentiated into
a larger female cell and a
smaller male cell ; in Vau-
rlii'i-in and Dichotomosiphon
oogamous reproduction is well
marked ( as ). The latter Algae
occur in fresh water or on damp
soil. The thallus consists of
a single branched, filamentous
cellattached to the substratum
by means of colourless rhizoids
(Fig. 296 Z>)-
The swarm-spores of Vau-
cheria, which differ from those
of the other Siphonales, are
developed in special sporangia,
cut off from the swollen ex-
tremities of lateral branches
by means of transverse walls
(Fig. 296). The whole con-
tents of such a sporangium be- FIG. 205. Caulerpa prollfen'. The shaded lines on the
tliiillus leaves indicate the currents of protoplasmic move-
come converted into a single,
ment ; a, growing apex of the thallus axis b, b, young
;

green swarm-spore. The wall thallus lobes ; r, rhizotds. (A nat. sizi>.)

of the sporangium then rup-
tures at the apex, and the swarm-spore, rotating on its longitudinal axis, forces
its way through the opening. The swarm-spore is so large as to be visible to
the naked eye, and contains numerous nuclei embedded in a peripheral layer
of colourless protoplasm. It is entirely surrounded with a fringe of cilia, which

protrude in pairs, one pair opposite each nucleus. Morphologically the swarm-
spores of Vnuchcria correspond to the collective, individual zoospores of an ordinary
sporangium.
The sexual reproduction of Vaucheria is not effected, like that of the other
Siphoneae, by the conjugation of motile gametes, from which, however, as the
earlier form of reproduction, it may be considered to have been derived. The
oogonia and antheridia first appear as small protuberances, which -grow out into
short lateral branches, and become separated by means of septa from the rest of
the thallus (Fig. 297 o, a). At first, according to OLTMANNS, the rudiment of an
362 BOTANY

Fi<;. L'iKi. \\i\K-li' fit- >r,s.v(/

.1. Yolinj: sporangium,
('.-. /.'. /onsporc with the s|>oran.uium from wliieh

it lias
escaped. ('. A pint ion of the peripheral /one of a zoos|x>re. />. a voiin^ plant with

rhizoids develo]ied from a xoos|Kire. (X. /'. after COT/.: /'. after S.\< us from OLTMANNS' :

Aline. I', after STI:ASHCKCKK.)

oogoniuni contains nuineroiis nuclei,

of which all hut one, the nucleus of
the future egg-cell, retreat again into
the main filament before the format ion
of the septum. In l'.<ji',,t inafd. 1>.\\ is
foun<l that all the nuclei, except that
of the oosphere, that arc in the
oogoninm degenerated and
disap-
peared. In its mature condition the
oogoniuni lias on one side a lieak-likc
projection containing only colourless
Fie:. li'.'T. I'liin-li'-i-iii .wxsi'/i.v. 1'ortion of a lilaineii
protoplasm. The oogoniuni opens at
with an oogoniuni, o antheruliiini, <i <-h, chro
this place, the oosphere rounding it-
; ;

iiiato]ili<>res : ii. cell nuclei; ut, nil plol'iiles.

self off. The antheridia, which are also
(X 240.)
multinuclear, are more or less coiled
(a), and open at the tip to set free their mucilaginous contents, which luvak up into
SECT. 1 CRYPTOGAMS 363

a number of swarming spermatozoids. The spermatozoids are very small, and have
a single nucleus and two cilia inserted on one side. They collect around the
receptive spot of the egg-cell, into which one spermatozoid finally penetrates.
After the egg-cell has been fertilised by the fusion of its nucleus with that of
the spermatozoid, it becomes invested with a wall and converted into a resting

oospore. On germination the oospore grows into a filamentous thallus.

CLASS X
6>36
Characeae (Stoneworts) ( )

The Characeae form a sharply defined group of Thallophytes,

the origin of which must be looked for in the Chlorophyceae, from
which, however, they are distinguished
by the complicated structure of their
sexual organs. They grow in fresh or
brackish water, 'attached to the bottom
and covering extended areas with a mass
of vegetation. In some species their
cylindrical main axes are over a foot in
length, and are composed of long inter-
nodes alternating with short nodes, from
which short, cylindrical branches are given
oft' in regular whorls with a similar
structure, but of limited growth (Fig.
298). The lateral axes are either un-
branched or give rise at their nodes to
outgrowths of a second order.
verticillate
From the axil of one of the side branches
of each whorl a lateral axis resembling
the main axis is produced. The attach-
ment to the substratum is effected by
means of branched rhizoids -springing
from the nodes at the base of the axes.
Both the main and lateral axes grow
in length by means of an apical cell, from
which other cells are successively cut oft'
by the formation transverse walls.
of
Each of these cells again divided by a
is

transverse wall into two cells, from the

lower of which a long, internodal cell
Flo. 298. Chara fragttis. End of
develops without further division while
; main shoot. (Nat. size.)
the upper, by continued division, gives
rise to a disc of nodal cells, the lateral axes, and also, in the lower

portion of the main axis, to the rhizoids. In the genus Nitella the
long internodes remain naked, but in the genus Chara they become
364 BOTANY

enveloped by a cortical layer consisting of longitudinal rows of

cellswhich develop at the nodes from the basal cells of the lateral
axes.
As a result of the fragmentation of its original nucleus, each inter-
nodal cell is provided with a number of nuclei which lie embedded

in an and actively moving layer of parietal protoplasm.

inner
Numerous chloroplasts are found in the internodal cells.
Asexual reproduction by means of swarm-spores or other spores

Fio. 299. A, Median longitudinal section through a lateral axis r, and the sexual
Chara fragilis.
organs which bears (x 60); a, antheridium borne on the basal nodal celllna, by the stalk
it

cell p; m, manubrium ob, an oogonium ; no, nodal cell ; po, the stalk-cell ; v, pivotal cell ;
;

c, (the crown B a lateral axis (bearing axes of the third order (x 6); a, antheridium; o,
oogonium.

is
unrepresented in the Characeae. Sexual reproduction, on the other
hand, is provided for by the production of egg-cells and spermatozoids.
The female organs are ovate. They are visible to the naked eye,
and, like the spherical red-coloured antheridia, are inserted on the
nodes of the lateral axes. With the exception of a few dioecious
species, the Characeae are monoecious.

The oogonia (Fig. 299 ob) contain a large ovum, filled with starch grains and
oil-drops ;
this is surrounded by spirally wound tubular cells forming the envelope.
These tubes terminate in the crown (c) between slits in which the spermatozoids
enter. The antheridia (Fig. 299 a) possess a wall formed of eight flat cells with
 CRYPTOGAMS 365

infolded walls (shields). From the centre of each of the shields a large cylindrical
cell(manubrium) projects into the central cavity, and the inner end of this bears
a number of head -cells. The latter bear long septate filaments from each cell
of which a spirally wound biciliate spermatozoid is formed. The form of the
spennatozoid (Fig. 98 A) differs considerably from that of other Algae.
The
egg. after fertilisation, now converted into an oospore, becomes invested
with a thick, colourless wall. The inner walls of the tubes become thickened
and encrusted with a deposit of calcium carbonate, while the external walls of
the tubes soon become disintegrated.
In a few cases, e.g. Chara crinita, the ovum can develop parthenogenetically
without being fertilised. Only female examples of this plant occur in Europe.
The oospore, on germination, gives rise first to a simple, filamentous row of
cells, the proem bryo. From the first node of the proembryo rhizoids are produced,
while at the second node there arise, together with a few simple lateral axes, one
or more main axes, which finally develop into a full-grown plant.
The formation of tuber-like bodies (bulbils, starch-stars) on the lower part of
the axes is characteristic of some species of the Characeae. These tubers, which
are densely filled with starch and serve as hibernating organs of vegetative repro-
duction, are either modified nodes with much -shortened branch whorls (e.g. in
Tolypellopsis stelligcra, when they are star -shaped), or correspond to modified
rhizoids (e.g. the bulbils of Chara ospera).

CLASS XI

20 37
'

Phaeophyceae (Brown Algae) (> )

The Phaeophyceae, like the Chlorophyceae, can be derived from

unicellular Flagellata, and from the Chrysomonadinae
in particular
which possess yellow chromatophores. On to these may be connected
unicellular colonial forms like Phaeococcus, or forms like PhaeotJiamnioit
composed of a short filament. The reproduction of these by means
of zoosporesand conjugating gametes already agrees with that of
the simpler Brown Algae.
With the exception of a very few fresh -water species, the
Phaeophyceae are only found in salt water. They are all attached,
and attain their highest development in the colder waters of the
ocean. They show great diversity in the form and structure of their
vegetative body. The simplest representatives of this class (e.g. the
genus Edocarpus) have a filamentous thallus consisting of a branched
or unbranched row of simple cells. Some Phaeophyceae, again, have
$. cylindrical, copiously branched, multicellular thallus (e.g. Cladotttephvs,
whose main axes are thickly beset with short multicellular branches
(Fig. 7) ;
while in other cases the multicellular thallus
is
ribbon-shaped
and dichotomously branched Didyota, Fig. 8).
(e.g.
Growth in length
in both of these forms ensues from the division of a large apical cell

(Figs. 7 and 132). Other species, again, are characterised by disc-

shaped or globose thalli.
366 BOTANY

The Laminariaecae and Fucaceae include the most highly developed

forms of the Phaeophyceae. To the first family belongs the genus
Laminaria found in the seas of northern latitudes. The large-
stalked thallus of the Laminarias resembles an immense leaf it is ;

attached to the substratum by means of branched, root-like holdfasts,

developed from the base of the stalk.

In Laminaria diyilata and L. Clonstoni (Fig. 301), a zone at the base of

Fio. 300. Macrocystis'pyrijem, Ag. ; a,

younger 6, older thallus. (A ft IT
SKOTTSBERO. ^, nat. size.)

the palmately divided, leaf -like expansion

of the thallus retains its meristematic
character, and by its intercalary growth
produces in autumn and winter a succes-
sion of new laminae. The older lamina
becomes pushed up and gradually dies, while
a new one takes its place and becomes in turn
palmately divided by longitudinal slits. The
large size of their thalli is also charact< Nti<- !

of the Laminarias ; L. saccharina (North Sea),

for instance, is frequently 3 m. long and the
stalk more than 1 cm. thick.
The
greatest dimensions attained by any
of the Phaeophyceae are exhibited by certain
of the Antarctic Laminariaceae. Of these,
FIG. 301. Laminaria
Macrocystis pyrifera (Fig. 300) is noted for its Sea. (Reduced to J.)
gigantic size ; the thallus grows attached to
the sea-bottom at a depth of 2-25 m., and, according to SKOTTSBERG ( M ), is at first
dichotomously branched. Single shoots of the thallus grow to the surface of the
water, and floating there attain a length of 70 m. they bear on one side long flat
;

lobes divided at their free ends, and having at the base of each a large swimming
bladder. Other noticeable forms, on account of their tree-like character, an- the
Antarctic species of Lessonia, in which the main axis is as thick as a man's arm ;
from it are given off lateral branches with hanging leaf-like segments. The plant
attains a height of several metres, and has a tree-like habit of growth.
 CRYPTOGAMS 367

The Fucaceae, although relatively large, do not compare with the Larninari-
aceae in size. As examples of well-known forms of this order may be cited Fucus
vesiculosus, which has a ribbon-shaped, dichotomously branching thallus with air-

FIG. 302. Fucus serratus. To the left the end of an older branch bearing conceptacles. (J nat. size.)

bladders, Fucus platycarpus without bladders, and Fucus serratus (Fig. 302). They
are fastened to the substratum by discoid holdfasts, and growing sometimes over
1 metre long, are found covering extended areas of the littoral region of the sea-

coast. Sargassum, a related genus chiefly inhabiting tropical oceans, surpasses

368 BOTANY

the other Brown Seaweeds, and even all other Algae, in the segmentation of its

thallus. The thallus of Sargassum shows', in fact, a distinction into slender,

branched, cylindrical axes with lateral outgrowths, which, according to their

function, are differentiated as foliage, bracteal, or fertile segments, or as air-

bladders. Various species of Sargassum which have been swept away from the
coast by currents finally collect in large floating masses in quiet regions of the
ocean (Sargasso Sea).

The cells of the Phaeophyceae have usually but one nucleus. They
contain a larger or smaller number of simple or lobed, disc-shaped
chromatophores, which contain a brown pigment (phaeophyll, p.
62), -giving to the algae a yellowish-brown or dark brown colour.
Numerous grains of a semi-fluid substance, to which the name
fucosan has been given, appear as the product of assimilation.
This substance appears to be a carbohydrate, and, according to
HANSTEEN ( 39 ), originates in the chromatophores. Among the more
highly developed forms the thallus exhibits a highly differentiated
anatomical structure. The quter cell layers, as a rule, function as
an assimilatory tissue, the inner cells as storage reservoirs. In
some species the axial cells of the thallus are arranged in strands
40
containing sieve-tubes ( ).
Two orders of Phaeophyceae may be distinguished. The Phaeo-
sporeae are vegetatively reproduced by means of zoospores and
sexually by ciliated gametes ; they thus resemble Ulothrix among the
Green Algae. The Cyclosporeae show a
marked differentiation of their sexual cells
into large, naked, non-motile, spherical
oospheres and small, ciliated spermatozoids;
some forms are also asexually reproduced by
means of naked, non-motile spores.

41
Order lt .
Phaeosporeae ( )

In this order are included the Laminarias, as well

as the majority of the other Phaeophyceae. Asexual
multiplication is effected by means of swarm-spores,

which are produced in large numbers in simple,

so-called unilocular sporangia they have a red eye-
;

spot, a chromatophore, and two LATERALLY ix-

si.i:n:i) cilia (Fig. 303J, one directed forwards and

the other backwards. Only this asexual reproduc-

tion yet known in the Laminariaceae.
is
FIG. 303. A, Pleuroeladia lacmtris. In addition to unilocular sporangia, multilocular
Unilocular sporangium with its
contents divided up into the sporangia are produced in the Phaeosporeae (Fig.
zoospores ; a, eye - spot ; ehr, 304). Each cell of the latter produces a single
chromatophore. .(After KLE- swarm -spore, rarely several. The conjugation of
BAHN.) }}, Chorda Jttum. Zoo- these swarm-spores has been observed in some genera.
spores. (After REINKE.) (From Qn tw account these swarm . spores mus t be termed
OLTMANNS Algae.)
gametes, and the corresponding sporangia gamet-
angia. The degree of sexual differentiation varies, and in some cases the swarm-
 CRYPTOGAMS 369

spores produced in multilocular sporangia can germinate without conjugating,

as was seen to occur in Ulothrix among the Chlorophyceae.
Ectocarpus siliculosus (Fig. 305) 'will serve as an example of the mode of con-
jugation of gametes produced from multi-
locular gametangia. The gametes are
similar in form, but their different be-
IW9 II ijgifev-- -i

J i

y/ Nlllflsl'^
haviour allows of their distinction into
male and female which are formed in dis-
"
* \
VF: W
:

< I
tinct gametangia, borne on the same or
The female gamete be-
I Y* u
:
^ ' different plants.
comes attached to a substratum, and
numerous male gametes gather around
it (Fig. 305, 1). Ultimately a male
gamete fuses with the female to form a
zygote (Fig. 305, 2-9). This contains
after the fusion a single nucleus, but two

Fio. 305. Ectocarpttii sH-iculonu*. 1, Female

Flo. 304. Pluri- surrounded by a number of male
.uaniete
A, Ectucarpustsiliculosus.
locular sporangium liberating its con- gametes seen from the side. 3-~>, Stages in
;

tents (after THURET). the fusion of gametes. 6, Zygote after 24

S, C, D, Sphace-
laria cirrhosa, development of the pluri- hours. 7-9, Fusion of the nuclei in conjuga-
locular sporangium (after REISKE). tion, as seen in fixed and .stained material.

(From OLTMANNS' Alyue.) (/-.'). after BERTHOLD ; 6-9, after OLTMANNS.)

chromatophores, and soon becomes attached and surrounded by a cell-wall ; it

grows into a new plant.

In other Phaeophyceae the distinction between the two kinds of gametes is
expressed in their shape and size. The Cutleriaceae afford a particularly good
transition from isogamy to oogamy (*'-).
2 B
370 HOT A N V

Order 2. Cyelosporeae

Only a small number of forms belong to this

4:! '
Family 1.
Dictyotaceae ").(

family. Padina pavonia, which occurs in the Mediterranean, and Dictyota dichoi'niiu.
which is widely spread in the European seas (Fig. 8), are examples. They are
distinguished from the Fucaceae by bearing asexual and sexual organs on distinct
individuals. The spores are formed as in the Red Algae in sporangia usually ;

there are four spores (tetraspores), less commonly eight. They have no cell-walls
and are unprovided with cilia and must be termed aplanospores (Fig. 306, 1).
The oogonia and antheridia in Dictyota are grouped in son (Fig. 306, 2, 3) and

Fio. 306. Dictyota dichotoma. Tran>vi ]> sections of tin- thallns. 7. WHhtetruparaagim; .'.with
a group of oogonia 3, with a group of antheridia (after THUKET). I,, Si>ermatozoiil (;iftt-r
;

WILLIAMS). (From OLTMANNS' Algae.)

arise from adjacent cortical cells each of which divides into a stalk cell and the

oogonium (or antheridium). The peripheral cells of the antheridial group remain
sterile and form a kind of indusium. Each oogonium forms a single uninucleatc
oosphere, the antheridia become septate, resembling the plurilocular gametaujjia.
and each cell gives rise to a spermatozoid. This, in contrast to the spermatozoids
of other Brown Algae, has a single long ciliuni inserted laterally.
Dictyota is dioecious. The male and female plants
arise from the asexually

produced tetraspores ;
from the fertilised ovum
plants which bear tetraspores are
developed. In the tetrad division in the sporangia the number of chromosomes
becomes reduced from 32 to 16, and the reduced number is maintained in all the
nuclei of the sexual plants, the double number being again attained in fertilisation.
There is thus (in contrast to the condition of things in the Phaeosporeae) a true
alternation of generations. The sexual generation (gametophyte) and the asexual
 CRYPTOGAMS 371

generation (sporophyte) do not, however, show differences in structure as is the case

in the Bryophyta and Pteridophyta.
Family 2. Fucaceae (**). Asexual
reproduction is wanting in this order,
while sexual reproduction is distinctly
oogamous. The oogonia and antheridia
of Fueus are formed in special flask-
shaped depressions termed CONCEP-
TACLES,' which are crowded together
below the surface in the swollen
tips of the dichotomously branched
thallus (cf. F. serratus, Fig. 302).
The conceptacles of F. platycarpus
(Fig. 307) contain both oogonia and
antheridia, while F. vesiculosiis, ou
the contrary, is dioecious. From the
inner wall of the conceptacles, be-
tween the oogonia and antheridia,
spring numerous unbrauched, sterile
hairs or PARAPHYSES, of which some

protrude in tufts from the mouth of

the conceptacle (Fig. 307 p). The Flo. 307. Fueus platycarpus. Monoecious con-
antheridia are oval in shape, and are ceptacle with oogonia of different ages (o), and
clusters of antheridia (a) ; p. paraphyses. (After
formed in clusters on special short and
THURET, x circa 25.)
much-branched filaments (Figs. 307 a,
309, 1, 2). The contents of each antheridium separate into a large number of

f.
-
" -

mes

FIG. 308. A, Oogonium, the contents of which have divided into the eight egg-cells. B, OogonJam,
from which the contents (C) have been extruded. D, K, Liberation of the eight egg-cells, st,
st;ilk-cell mes, middle, and end, inner layers of the oogonial cell-wall. (After THURET. From
;

OI.TM ANNS' A!'j"<.)

spermatoxoids, which are discharged in a mass, still enclosed within the inner

layer of the antheridial wall (Fig. 309, 4). Eventually set free from this outer
372 BOTANY

covering, the spermatozoids appear as somewhat elongated, ovate bodies, having

two lateral cilia of unequal length and a red eye-spot. The oogonia (Figs. 307 o,
308) are nearly spherical, and are borne on a short stalk consisting of a single cell.
They are of a yellowish-brown colour, and enclose eight spherical egg-cells which
are formed by the division of the oogonium mother cell. The eggs are enclosed
within a thin membrane when ejected from the oogonium (Fig. 308 B, C). This
membranous envelope deliquesces at one end and, turning partly inside out, sets
free the eggs (D, E). The spermatozoids then gather round the eggs in such

FIG. 309. Fucus. 1, Group of antheridiu. ..'. 1'ait of an antliri idimii sliouin-
spermatozoid.s. "3, Spermatozokl o, eye-spot k, nucleus.
: ; ^ Isolated 'antheridia liberating tbfl
spermatozoids. 5, Oospliere surrounded by spermatozoids. 6, Section through a 1'<>i tili.-''<l i-.^.n :

ek, nucleus of epp : >',/>/.. sperm-nucleus ; sp, s))ermatozoids. (1, If, f>, after THURET ; 2, :>, a ft IT
GCIGNARD ; 6, after FARMER. From OLTMANNS' Algae.)

numbers that by the energy of their movements they often set them in rotation
(Fig. 309, 5). After an egg has been fertilised by the entrance of one of the
spermatozoids (Fig. 309, 6) it becomes invested with a cell-wall, attaches itself
to the substratum, and gives rise by division to a new plant.
In the case of other Fucaceae which produce four, two, or even only one egg in
their oogonia, the nucleus of each oogonium, according to OLTMANXS, nevertheless
first divides into eight daughter nuclei, of which, however, only the proper
number give rise to eggs capable of undergoing fertilisation.
Since the Fucaceae have no asexual spore- formation the alternation of genera-
tions characteristic of Dictyota is wanting in them. It is possible to regard the
 CRYPTOGAMS 373

divisions leading to the development of the sexual cells in the oogonium and
antheridium, in which the reduction of chromosomes takes place, as corresponding
to a reduced sexual generation. On this view the thallus of Fucus would corre-
spond to the sporophyte of Dictyota, and the young sexual organs would be
41
equivalent to the tetrasporangia of the latter ( ).
Economic Uses. The dried stalks of Laminaria digitata and L. Cloustoni are
used as dilating agents in surgery. IODINE is obtained from the ash (varec, kelp)
of various Laminariaceae and Fucaceae, and formerly soda was similarly obtained.
Many Laminarias are rich in MANNITE (e.g. Laminaria saccharina), and are used in
its production, and also as an article of food by the Chinese and Japanese.

CLASS XII

6> 20> 40
Rhodophyceae (Red Algae) ( )

The Rhodophyceae or Florideae constitute an independent group

of the higher Algae, the phylogenetic origin of which is perhaps to

FIG. 310. Chondrus traits. (^ nat. siw.)

be sought among the higher Green Algae. They are almost ex-
clusively marine, and specially characterise the lowest algal region
on the coasts of all oceans, especially in temperate and tropical
2 B 1
374 BOTANY

latitudes. A few genera (e.g. Batrachospermum, Lemanea) grow in

fresh-water streams.
Red Algae exhibits a great variety of forms.
The thallus of the
The simplest forms are represented by branched filaments consist-
ing of single rows of cells (e.g. Callithamnion). In other cases the
branched filamentous thallus appears multicellular in cross-section.
In many other forms the thallus is flattened and ribbon-like (e.g.
Chondrus crispus, Fig. 310; Gigartina mammillosa, Fig. 311); while
in other species it consists of expanded cell surfaces attached to a
substratum.
One of the more complicated forms is Delesseria sanguined
(Hydrolapathum) (Fig. 9), which occurs on the coasts of the Atlantic.
The leaf -like thallus which
springs from an attaching disc
is
provided with mid ribs and
lateral ribs. In the autumn the
wing-like expansions of the thallus
are lost, but the main ribs persist
and give rise to new branches in
the succeeding spring. All the
Florideae are attached at the base
by means of rhizoidal filaments
or discoid holdfasts. The thalli
of the Corallinaceae, which have
the form of branched filaments
or of flattened or tuberculate in-
crustations, are especially char-
acterised by their coral-like ap-
pearance, owing to the large
amount of calcium carbonate
FIG. 311. Gigartina mammillosa. s, Wait->haped
deposited in their cell walls.
-
cystocarps. (} nat. si/r.)
The calcareous Florideae are
chiefly found on coasts exposed to a strong surf, especially in the
tropics.
The Rhodophyceae are usually red or violet ; sometimes, however,
they have a dark purple or reddish-brown colour. Their chromato-
phores, which are flat, discoid, oval, or irregular-shaped bodies and
closelycrowded together in large numbers in the cells, contain a red
pigment, PHYCOERYTHRIN, which completely masks the chlorophyll
and appears to be chemically combined with it (p. 62). True starch is
never formed as a product of assimilation, its place being taken by
other substances, very frequently, for example, by Floridean starch,
in the form of spherical stratified grains which stain red with iodine.

Oil-drops also occur. The cells may contain one or several nuclei.
Reproduction is effected either asexually by means of spores, or
sexually by the fertilisation of female organs by male cells.
 CRYPTOGAMS 375

The asexual SPOKES are noil-motile ; they have no cilia and are simply naked,

spherical cells. They are produced, usually, in groups of four, by the division
of a mother cell or sporangium, from which they are in time set free by the
transverse rupture of its walls. The spor-
angia themselves are nearly spherical or
oval bodies seated on the thalloid filaments
or embedded in the thallus. The spores
escape by a transverse rupture of the wall
'

of the sporangium. In consequence of

their usual formation in fours, the spores
of the Florideae are termed TETRASPORES
(Fig. 312). They are analogous to the
swarm-spores of other Algae similar spores
;

are found also in the Dictyotaceae among

the brown Algae.
In the construction of the sexual
organs, particularly the female, the Rhodo-
phyceae differ widely from the other Algae.
Hatrachospermum moniliforme, a fresh-
FIG. 312. Callithamnionwrymbosii.ni. .4, Closed
water form, may serve as an example to
sporangium ; I), empty sporangium with
illustrate the mode of their formation. This four extruded tetraspores. (After THTTRF.T. )
Alga possesses a brownish thallus, en-
veloped in mucilage, and consisting of verticillately branched filaments. The

FIG. 313. Batrachospermum iiwniliformc. A, Male branch with antheridia, isolated 1>\ pressure ;

s*, a spermatium ;a .spermatium escaping from an antheridum r, an empty antheridium.

s, ;

J5, female branch wjtli an unfertilised carpogonium c, basal portion

; t, trichogyne of carjK)-
;

gonium. C, female branch with fertilised carpogonium ; s, the spermatium after the fusion of
its contents with the trichogyne ; c, fertile filaments developing from the basal portion of the
carpogonium. (x 540.)

sexual organs appear in the autumn and form on the branching whorls glomeruli
or spherical bodies composed of short, radiating branches.
The antheridia, also known as spermatangia (Fig. 313 A), are produced, usually
in pairs, at the ends of the radiating branches of a glomerulus. Each antlu-ridium
2 B 2
37 BOTANY i'\KT 11

consists of a single thin-walled cell, in which tin- whole of the protoplasm. ,i> is
tin- rule in all Khodophyceae, is consumed in the formation of <m<- uninuclear

si'KK.M.viiJ. M :
iii-cordiii^ to sonic authors the nucleus of the spcrmatium divides
iuto two. Tin- nearly spherical, and are invested with a thin
s|ierniatia an-
outer inenihi-iinc or cell wall. They are non-motile, unlike the ciliated sp< T-
niatozoids of the other Algne. anil have therefore received a distinctive name. In

consequence of their incapacity for independent movement, they must lie carried
passively by tlie water to the female organs, which arc situated near the
autheridia ;it the ends of other branches. The female organ is called a AK- <

POGONIUM (Fig. 313 .#), and consists of an elongated cell with a liasal, llask-
sha|ied portion (c) prolonged into a filament, termed the TI:H-HII;YXK (/). Tin-
basal portion contains the nucleus of the egg and the chromatophorcs. while tin-
trichogyne functions as a receptive organ for the spermatia, one or two of which
fuse with it. and the contents, escaping through the .spermathun wall, pass into
the nrpogonium. The sjK-rm nucleus juisses down the trichogyne and fuses with
the nucleus of the egg-cell. The fertilised egg, which becomes limited from
the trichogyne by a wall, does not become converted directly into an oospore.
but, as a result of fertilisation, numerous branching sporogenous filaments
grow out from the sides of the ventral portion of the carpogonium. At the same
time, by the development of outgrowths from cells at the base of tin- carpo-
goniuni an envelope is formed about the sporogenous filaments. The whole
product of fertilisation, including the surrounding envelope, constitutes the
fructification, and is termed a YSTIM'AKP. The profusely-branched sporogcnous
i

filaments become swollen at the tips and give rise to spherical, uninuclear s|Mire>
known as CAUPOSPOKES, which are eventually set free from the envelop.. In
the rase of Batrachospermum the carpos pores produce a filamentous protonema.
the terminal cells of which give rise to asexual unicellular spores. These spores
serve only for the multiplication of the protonema. Ultimately, however, one
of the lateral branches of the protonema develops into the sexually differentiated
filamentous thallus. The production of spores by the protonema is analogous to
the formation of tetraspon-s by other Florideae.
The formation of the cystocarps and carpospmvs j s much more complicated
in the case of other genera, but in all eases, according to
OI.TMAXXS, the carpooporea
are descended from the fertilised egg-cell. There arethus two generations dis-
tinguishable in the life-history of the Florideae, the sexual (gametophyte), which
bears the egg-cells and the spermatia. and the asexual generation sporophvte i
.

derived from the fertilised egg-cell the latter generation, which produces tin-
;

carposj>ores, remains in connection with the parent plant. This type of alter-
nation of generations is comparable with that found in Mosses and Ferns. The
production of tetraspores is an asexual mode of reproduction of the sexual genera-
tionand precedes the formation of the sexual organs.
Diulresnaya coccinen, which is found on the warmer coasts of Europe, has a
branched, cylindrical thallus and will serve as an example of the more complicated
mode of origin of the spore-bearing generation (Fig. 314). The carpogonial
branches consist of about seven cells, the terminal one bearing a very long
trichogyne. After fertilisation the carpogonial cell grows out into a filament,
which elongates and becomes branched. This filament fuses with a number of
special cells, characterised by their abundant contents, the u XII.IAKY CKI.I.S. Tin-
first of these lie in the carpogonial branch itself, the others in adjoining lateral

branches. All the nuclei of the sporogenous filament are derived by division from
that of the fertilised egg-cell. The successive fusions with auxiliary cells do not
 CRYPTOGAMS 377

involve nuclear fusions, but simply serve to nourish the sporogcnous filament.
A second and third sporogenous filament may arise from the carpogonial eell.

B
KKI. :;i4. Dttdretnaya <-<i<-i'inii<. .1 Cari>o;;oiiiai i>ranch; a, carpogoniom ; t. trichoxyne. 7.'. car|M>-
xonium after fi>i-tilis;itii)ii. into tin- siwinixfiious liUiiiicnt (.-/).
^rowu (int <'. fusion of tin-
.-IHiro^i'iioiis lilanirnt with the first :mxilinry cfll ('(). /), liranchinK of the lilainent and fusion

with six auxiliary ((Us ("i-",;); the cells ;-,; ai'e borne on bnnche* qrigllUttillg from tin- axis
ha (diagrammatic). /,'. Ki]M- cluster of cai-ix>s]Mires 01 ij;inatiu^ from one branch. (.!-/>. nlXei-

Ol.TMANNs: A', alter HoKNKT. A-<', >' alHlllt.'iOO; />. Y. 250 E, X 300.) :

Two outgrowths now arise from each of the swollen cells of the sporogeiious
filament which fused with auxiliary cells. By further division oT these outgrowth.--
378 BOTANY VAI:T n

the spherical masses of carpospores, which subsequently become free, are derived
(Fig.314 E).
47
Harveyella mirabilis ( ), one of the Florideae occurring in the North Sea, is of
special interest. It grows as a parasite on another ivd .seaweed, RhodomeJu xuli-

fusca, on which it appears in the form of a small white cushion-like growth.

As a result of its parasitic mode of life the formation of chromatophoivs Las n 1

entirely suppressed, and, thus this plant behaves like a true Fungus.
Economic Uses. Giyartina mammillosa (Fig. 311), with peg-like cystocarps
2-5 mm. in length, and Chondrus crispus (Fig. 310), with oval cystocarps about
2 mm. long, sunk in the thallus, tetraspores in groups on the terminal segments
of the thallus. Both forms occur in the North Sea as purplish-red or purplish-
brown Algae ;
when dried they have a light yellow colour, and furnish the official
CARRAGHEEN, "Irish Moss," used in the preparation of jelly. AGAR-AGAR,
which isused for a similar purpose, is obtained from various Florideae Gmciluria :

lichenoidts supplies the Agar of Ceylon (also called Fucus amylaceus), Eucheuina
spinosum, the Agar of Java and Madagascar.

CLASS XIII

4S> 49> 50
Phyeomycetes ( )

In the nature both of their thallus and sexual organs the Phyeo-
mycetes exhibit a close connection with the Siphoneae. The phylo-
genetic origin of most of the Phyeomycetes must be sought in this
group, though certain forms point to a relationship with other (In-m
Algae (e.g. Basidiobolus with the Conjugatae). They can only for the
present be regarded as a definite class, pending their separation into
several series derived from distinct classes of Algae.
The thallus consists of extensively branched tubular threads in
which, as is the case in Vaucheria, transverse septa only form in
connection with the reproductive organs. The continuous proto-
plasmic mass contains numbers of very small nuclei, but chromato-
phores are entirely wanting in these colourless organisms. The
whole thallus a Fungus is
of spoken of as the mycelium, the
individual filaments as hyphae. In the Phyeomycetes the hyph;i-
are non-septate, their division into distinct cells only taking place
in a few cases. The plants are either saprophytes occurring on the
putrefying remains of animals or plants in water or on decaying
organic substances exposed to air, or they live parasitically in the
tissues of higher plants or on insects.
Asexual reproduction is effected by means of spores. These are
formed in the majority of the genera within sporangia, the proto-
plasm of which splits into the numerous spores. The latter escape
in the genera which live in water as ciliated swarm-spores (Fig. 316) ;

in the forms which are exposed to the air the spores are enclosed in
a cell wall (Fig. 323). The conidia, which are sometimes found
 CRYPTOGAMS 379

together with sporangia, in other cases alone, are also adapted for
dispersal inair. They arise by a process of budding and abstriction
from the ends of certain hyphae which are usually raised above the
substratum as special conidiophores.
The sexual organs of the Phy corny cetes are in many ways
peculiar, and the two groups of the Oomycetes and the Zygomycetes
are distinguished according to their nature. In the Oomycetes,
which stand nearer to the Green Algae, oogonia and antheridia are
found the contents of the latter enter the oogonium by means of a
;

tubular outgrowth, and after fertilisation oospores are formed. In

Monoblepharis alone are free spermatozoids found. The sexual
organs of the Zygomycetes are alike, and on conjugation a zygospore
is
produced. They are usually multinucleate and thus are morpho-
logically comparable to a whole gametangium of an isogamous Alga.
Multinucleate gametangia, oogonia, and antheridia, which fuse
directly with one another, without the separation and escape of the
individual gametes, are generally termed CCENOGAMETES.

Order 1. Oomycetes
51
1. Only in the small family of the Monoblepharideae ( ) are free ciliated

s]n-nii:itozoids liberated from the antheridia. In the other Oomycetes the multi-
nucleate contents of the antheridium do not divide into separate spermatozoids,

Pio. 315. Monoblepharis sphaerica. End of filament with terminal oogonium (o) and an antheridium
(a) : before the formation of the egg-cells and spermatozoids 2, spermatozoids (s) escaping
1, ;

and approaching the opening of the oogonium 3, osp, ripe oospore, and an empty antlieridinm.
;

( After CORNU, x 800.)

but are directly introduced into the egg-cell by means of an outgrowth of the
antheridium.
The species of Monoblepharis live in water upon decaying remains of plants.
Asexual reproduction is effected by means of swarm - spores, formed in large
numbers in sporangia. The oogonium, which is usually terminal, contains only a
380 BOTANY

single egg-cell (Fig. 315). The antheridia, which resemble the sporangia, liberate
a number of uniciliate spermatozoids. On a spermatozoid reaching the egg-cell
through an opening in the tip of the oogonium an oospore is formed. A spinous
cell wall forms around the oospore.
52
2. The Saprolegniaceae ( ), which connect on to the preceding family, live like
them saprophytically on the surface of decaying plants and insects ami even on
living fishes. Asexual propaga-
tion is effected by club-shaped .

sporangia (Fig. 316) which pro-

duce numerous biciliate swarm -

spores. The sexual organs develop

Q
on older branches of the mycelium
(Fig. 317 a). The oogonia give
rise to a larger (as many as 50)
or smaller number
of egg-cells,

rarely only to a single one. At

first the egg-cell contains numer-

Fio. 317. Saprolegnia mixta. Hyphre bearing

the sexual organs ; o, antheridium which has
sent a fertilisation tube into the oogonium ; o',
FlO. 816. Saprolegnia miita. The egg-cell ; o", oospore enclosed in a cell-wall ; op,
biciliate zoospores, s2 ,
are es parthogenetic oospores ; g, young oogonium.
from the sporangium. (After G. KLEBS.)

ous nuclei, but usually all but the single nucleus of the egg degenerate. The
tubular, multinucleate antheridia apply themselves to the oogonia and send
fertilising tubes to the egg-cells. One male nucleus enters the egg-cell and
fuses with its nucleus (Fig. 318). The oospore after fertilisation acquires a
thick wall. In some forms belonging to this family and to the Peronosporeae
the formation of antheridia is occasionally or constantly suppressed the oospores ;

develop parthenogenetically without being fertilised (Fig. 317).

3. The Chytridiaceae ( M ) are microscopically small Fungi parasitic on aquatic or
 CRYPTOGAMS 381

land plants and in some cases on animals. The non-septate mycelium is feebly
developed, and is frequently reduced to a single sac-shaped cell inhabiting a cell
of the host. Asexual multiplication is effected by means of swarm-spores provided
with one or two cilia. In the simplest forms the entire cell becomes converted
into the sporangium. Olpidiuni Brassicae (Fig. 319), which is parasitic in the
tissues of the base of the stem of young

cabbage plants and brings about their

death, may be taken as an example. The
sporangia in this case have a long neck
from which the swarm-spores escape.
Thick-walled sporangia which undergo
a resting period are also frequently found
in the Cliytridiaceae.
Sexual reproduction is only found
in some of the genera, in the form of
fusion of an antheridium with an oogou-
"
ium and the production of an oospore. Fia 31S.-Acki^ i,ol,,,, nd,-a.
. Th,- fertilisation of
The systematic position of the Chytri- two egg-cells, o, of an oogonimn by two tubes
diaceae among the Phycomycetes is still from the antheridium a ek, nucleus of the egg-
;

doubtful the simplest forms appear to cell ;sperm-nucleus; in o* the s.-rtion has
sk,
;
not passed through the egg-nuelfiis. (After
have originated in part from Flagellatae
TROW.)
and partly from Protococcaceae.
4. The Peronosporeae (
r>4
) Fungi whose mycelium penetrates the
are parasitic
tissues of the higher plants, and is frequently the cause of death. In damp climates,
certain species occasion epidemic diseases in cultivated Thus, the mycelium
plants.
of Phytophtliora, infestans, the fungus which causes the Potato disease, lives in the
intercellular spaces of the leaves and tubers of the Potato plant, and
by penetrating
the cells with its short hau-
storia it leads to the discolora-
tion and death of the foliage
and Sexual reproduc-
tubers.
tive organs have not as yet
been observed in this species.
Asexual, oval sporangia are
formed on long branching
sporangiophores which grow
out of the stomata, particu-
larly from those on the under
side of the leaves (Fig. 320),
and appear to the naked eye
" as a white mould. The spor-
Fio. sw.ulpldium. Hrassim,: A, Thiv,- zoosporangia, th.'
angia, at first terminal, are cut
contents of one of which has escaped (x 100) B, Zoo- off b J transverse walls from
s p ores (x o-20). '. Kiting sporangia (X 520). (Aftpr
the ends of the branches of the
WOROXIX.)
sporangiophore, by the subse-
quent growth of which they become pushed to one side, and so appear to be inserted
laterally. Before any division of their contents has taken place, the sporangia (B)
fall off and are disseminated
by the wind in this way the epidemic becomes
;

widespread. The development

of swarm-spores in sporangia is effected only in
Water, and is consequently possible only in wet weather. In this process the
contents of the sporangium divide into several biciliate swarm-spores (C, D). Each
 BOTANY

of these spores after escaping from the sporangium gives rise to a mycelium, which
penetrates the tissues of a leaf. The sporangium may also germinate directly
without undergoing division and form ing swarm -spores. A similar transformation
of sporangia into conidia is also found in other Peronosporcae as a result of their
transition from an aquatic to a terrestrial mode of life.
rifii-ii/ii. an extremely destructive parasite, also produces copiously

branched sporangiophores and occa-

sions the "False Mildew" of the
leaves and fruit of the Grape-vine.
A Ibugo Candida = Cystojnis candidus),
(

another very common species, occurs

on 'rncil'erae. in particular on fiijixi-lfn
(

blttrmpcutoris, causing wliite swellings

on the stems. In tliis species the
sporangia are formed in long chains
on the branches of the mycelium
under the epidermis of the host plant.
and produce numerous .swarm-spores.
The sexual organs of the Perono-
sporeae resemble those of the genus
raucheriu. (Fig. 292). They arise
within the host plant the oogonia as
spherical swellings of the ends of
certain hyph.v, the antheridia as
tube-like outgrowths arising as a
rule just below the oogonia. Both
are cut off by transverse walls and
are multinucleate (Fig. 321). The
several species exhibit interesting
differences as regards the nuclear

changes. In Ptfonospora /i>/i-n.<///<-n,

Albuyu Candida, and A. Lcpigom, l'[i-

t/iiniii. rinfitiii/tni'ti. ami Sdtrospora,

a single large central egg-cell or

oosphere becomes differentiated in

the protoplasm of the oogoninm this ;

contains a single nucleus in a central

position, while the remaining nuclei
Ki<;. 3-JO.-.I. Surface vi.'u th- epi of a
pass j nto t l, e peripheral layer of pro-
,,f

Mntu 1,-i.f. with of fhntorhtho,-,'

t lasm (perf.^,,,). Theantheridium
ipc^opfaorai
in/Man* projecting t'rnni the atomata ( x (HI); ;;.
now sends a process into the oogonmm,
, ,i,,- iponngfam; C, anotl.-r in proceM of
division ; /), a swnnn-srxMH. which at its apex opens into the
(;,-/> x 540.)
oosphere and allows a single male
nucleus to pass into the latter (Fig. 321, 2). The oosphere then becomes surrounded
with a cell wall (Fig. 321, 3), and nuclear fusion takes place, while the periplasni
is membrane of the spore (episporium). In Perono-
utilised in forming the outer
the ripe oospore has a single nucleus, in Albuyo it becomes
sj>orn fiu ?< sit! i-i<
i

multinucleate as a result of nuclear division. In J/6 <// I'liti and A. pvrtulacac

there is also a central oosphere surrounded by periplasm, but the oosphere contains
numerous nuclei, which fuse in pairs with a number of male nuclei entering from
the anthoridium. A multinucleate oosporo thus arises from the compound egg-cell.
 CRYPTOGAMS 383

The behaviour of these two species can be regarded as primitive, the uninucleate
oospheres of the first -named forms having been derived from the multinucleate

condition. Albuijo tragopoyouis occupies an intermediate position in that its

oosphere at first multinucleate, but later contains only one female nucleus, tin-
is

others having degenerated. The superfluous nuclei in the oogonia and antheridia
may be regarded as the nuclei of gametes which have become functionless, and are
comparable with the superfluous egg-nuclei of certain Fucaceae. The oospores
either produce a mycelium directly or give rise to swarm-spores.

FIG. 321. Fertilisation of the Peronosporeae. 1, 1'eronospora

parasitica. Young multinucleate oogonium (og) and anther-
idiuin (an). %, Allunjo Candida. Oogonium with the central FIG. 322. Rhizopiis nigricans ( Mm-or sto-
uninucleate oosphere and the fertilising tul>r (</) of the lonifer). Portion of tin- niyceliuin with
anthcridiuni which introduces the male nucleus. 3, The sam>-. three sporangia that to the right is shed-
;

Fertilised egg-cell (o) suiTounded by the periplasm (/.). (After ding its sjwres and shows tint jiersisti-nt
\V MiK.K. x 666.) hemispherical columella. (x 38.)

Order 2. Zygomycetes

1. The Mucorineae (
55
) comprise a number of the most common Mould Fungi.
They are saprophytic, and are found chiefly on decaying vegetable and animal
substances. Asexual reproduction is effected by non-motile, walled spores, which
either have the form of conidia or arise endogenously in sporangia. Sexual
reproduction consists in the formation of zygospores, as a result of the conjugation
of two equivalent coenogametes.
One of the most widely distributed species is Mucor Mucedo, frequently found
forming white fur-like growths of mould on damp bread, preserved fruits, dung,
etc. Mucor stolonifcr (^lihizopus nigricans}, with a brown mycelium, occurs on
similar substrata. The spherical sporangia are borne on the ends of thick erect
branches of the mycelium (Fig. 322). From the apex of each sporangiophore a
.single spherical sporangium is cut off by a transverse wall, which protrudes into
384 BOTANY I'AKT II

the cavity of the sporangium aiid forms a columella (Fig. 323, 1, c). The contents
of the sporangium become divided into numerous spores. These escape by the
swelling of a substance which lies between the spores and the bursting of the
sporangial wall. In Pilobolus, which occurs commonly on dung, the sporangium
is forcibly cast off from the turgid sporangiophore which bursts at the columella.

Fio. 323. 1, Mucor Mwxdo.

Asporangium in optical longitudinal section <. ccilumella m. wall
: ;

of sporangium sp, spores. 2, Mwxtr mucilagineuf. A sporangium shedding its spores ; the
;

wall (i) is ruptured, and the mucilaginous substance between the spores (<) is greatly swollen.
(After BREFELD, 1 x 225, 2 x 300, from v. TAVKI.. /;/:,-.)

According to HARPER the spores of Pilobolus are binucleate, while those of Sporo-
dinia (Fig. 324) are multinucleate.
conditions, instead of asexual sporangia, organs of sexual repro-
Under certain
duction are produced. The hyphae of the mycelium then give rise to lateral, club-
shaped branches. When the tips of two such branches come into contact, a
conjugating cell or ccenogamete is cut off
from each by a transverse wall (Fig. 325).
The two gametes thereupon coalesce, their
nuclei conjugating in pairs, and fuse into
a ZYGOSPORE, the outer wall of which is
covered with warty protuberances. After
a period of rest the zygospore germinates,

developing a germ-tube, which may at

once bear a sporangium (Fig. 325, 5).
BLAKESLEE'S ( x ) demonstration of the
dioecious nature of the mycelium of Mucor
stolonifer is of great interest. The formation
of zygospores only takes place when male
Fi<;. 324. Sporwlinia grandif. Median section an(j female myce li a come j u contact. In
of a sporangium.
ripe The spores are
multinncleate. (After HARPER, x 4-25.)
Mucorineae the two conjugating
other
gametes may arise on the same mycelium.
Within the group of the Zygomycetes a reduction of sexuality is perceptible.
Thus, in the case of certain -Mucoriueae, although the conjugating hyplite meet in
pairs, no fusion takes place, and their terminal cells become converted directly
into spores, which are termed AZYGOSPORES. In other forms again, hyplm-
producing azygospores are developed, but remain solitary, and do not, as in the
preceding case, come into contact with similar hyphse. There arc also many
species in which the formation of zygospores is infrequent.
Both the size and number of spores produced in the sporangia of M-ttfnr
 CRYPTOGAMS 385

are subject to variation. The sporangia of the genus Thamnidium are, on the other
hand, regularly dimorphic, and a large sporangium containing many spores is formed
at the end of the main axis of the sporangiophore, while numerous small sporangia,
having but few spores (sporangioles), are produced by its verticillately branching
lateral axes. The sporangia may at times develop only a single spore, as the
result of certain conditions of food-supply, and in this way assume the character
of conidia. This dimorphism is
even more complete in the trop-
ical genus Choanephora. In this
case, in addition to large spor-
angia, conidia are produced on
special conidiophores. There are,

finally, Zygomycetes (e.g. Chaeto-

cladium) whose only asexual
spores are conidia. In this one
group, therefore, all transitional
forms, from many-spored spor-
angia to unicellular conidia, are
represented.

Flo. 3'25. Mm-oi- Mucrdo. Different .stages in the

formation and germination of the xyj^^pore.
1, Two conjugating branches in contact; ,.'.

septation <>f tlic conjugating cells () from the FIG. 32(i. Empusti Mvrai: A, Hypha
siispeiiM>r> (//) ; J, more advanced sta.^e. th. from the body of a fly. Ji, Young
conjugating cells (a) are still distinct from one conidiophore arising from the my-
another the warty thickenings of their walls
: celium and projecting from the
have commenced to form it, ripe xygospore (//)
; body of the insect. C, Formation
between tln j

snspeiisors (a)', germinating: '>. of the conidimn into which the

xygospore with a p rm-tnlie bearing a spor-
j
numerous nuclei have passed from
angium. (After BREFELD, 1-4 x 2'25, "> x circa the conidiophore. (After OLIVE,
60, from v. TAVKI., /'//:..) x 540.)

group of Fungi which mostly In v

57
2. The Entomophthorineae ( )
is a small

parasitically in the bodies of insects and caterpillars. The multinucleate mycelium

remains non-septate or later becomes divided into cells. Asexual multiplication is
effected by means of conidia which contain one or numerous nuclei. These arix;
singly at the ends of branches of the mycelium and when ripe are forcibly abjected.
Sexual reproduction is by means of zygospores, in place of which azygospores
frequently urisc.
2 c
386 BOTANY PART n

The best-known example is Empusa 'inuscae (Fig. 326), which is parasitic on

house-flies. The couidia, which are multinucleate, form a white halo around the
body of the dead fly which has by the fungus.
In-en killed
3. Basidiobolaceae ('*}.- ranarum, a saprophytic Fungus growing
-Jiasidiobolus
on the excrement of Frogs, must be separated from the preceding group. Each of
the cells of its septate mycelium contains one large nucleus. The conidia which
arise singly on the ends of the conidiophores and are abjected when ripe are
uninucleate. The mode of origin of the zygospores is peculiar. Two adjoining
cellsconjugate after they have put out beak-shapcil processes which are cut off as
transitory cells. In the zygospore the two sexual nuclei give rise to four, of which
two disorganise while the other two fuse. Both in this procedure and in the
nuclear structure there are evident resemblances to the Conjugatae (Spirogyra).

CLASS XIV
ls 41
Eumycetes ( ')

When the Phycomycetes are excluded there remain two great

groups of Fungi, the Ascomycetes and the Basidiomycetes, regarding
the classification and phylogeny of which' there is still much un-
certainty. The attempt has been made to derive them from the
Phycomycetes. Not only is the construction of the thallus against
this, but the structure of the sexual organs and the development of
the fruit in the Ascomycetes indicate on the other hand a remarkable
relationship with the Red Algae. The Uredineae or Rusts, one of
the simplest orders of Basidiomycetes, appear to connect the latter
group with the Ascomycetes.
The saprophyticor parasitic thallus of the Eumycetes is, like that
of the Phycomycetes, composed of fine, richly branched filaments or
hyphse which together form the mycelium. The hyphae are, how-
ever, in this case septate, consisting of rows of cells. The cell-
membrane, which contains chitin, is usually thin. In the colourless
protoplasm there are usually numerous minute nuclei (Fig. 62),
while in other cases each cell has a pair of nuclei or only a single
nucleus. Chromatophores are wanting and true starch is never
formed the place of the latter is taken by glycogen, often in con-
;

siderable quantity, and by fat-globules. The hyphse of a mycelium are,

as a rule, either isolated or only loosely interwoven ; they spread
through the substratum in all directions in their search for organic
nourishment. In many of the higher Fungi, however, the profusely

branching hyphae form compact masses of tissue, AVhere the fila-

ments in such cases are in intimate contact and divided into short
cells, an apparently parenchymatous tissue or PSEUDO-PARENCHYMA
is produced. Such compact masses of hyphal tissue are formed by
some species of Fungi when their mycelia, in passing into a vegetative
resting stage, become converted into SCLEROTIA, tuberous or strand-
SECT. I 387

like, firm, pseudo-parenchymatous bodies, which germinate under certain

conditions (Fig. 107). In the fructifications the hyphae are also nearly
always aggregated into a more or less compact tissue (Fig. 106).
The distinction between the two main groups is apparent in their
reproductive organs.
1. The Ascomycetes in their typical forms
possess sexual organs.
The oogonia, Avhich here go by the name of ascogonia or, as in the
Red Algae, of carpogonia, show profound differences in the genera
which have as yet been accurately inves-
tigated and the same may be said of the
male organs. In all cases, however, the
fertilised carpogonium does not become a

resting oospore, but, while in connection

with the maternal plant (gametophyte),
develops into an asexual generation (sporo- a
phyte). In this process hyphse bud out
from the carpogonium, and their ultimate
branches terminate in the asci which are
sporangia of a specialised type.
The ASCUS ( 60 ), which is so character-
istic of the whole
group, originates from a
tubular cell ; this to start with contains
two nuclei which fuse, and the resulting
nucleus by repeated division gives rise to
eight nuclei. By a process of free cell-
formation the spores become limited by
cell-walls in the way shown in Fig. 96

(Figs. 327, 334). In contrast to the forma- FIG. 327. Portion of the hyim-iimni
tion of spores in the sporangia of Phyco- of Morchella esculentu. a, Asci ;

p, paraphyses sh, Milihynu-iiial

;

mycetes the cytoplasm of the ascus is not tissue, (x 240.)

completely used up in the formation of
the ascospores. The spores usually form a longitudinal row, and
are ultimately ejected from the ruptured apex of the ascus by the
swelling of the remaining cytoplasm. In the great majority of the
Ascomycetes the number of ascospores in the ascus is eight.
In the majority of Ascomycetes the asci, which originate from the
carpogonium, are associated in special fructifications vegetative ;

filaments of the mycelium take part in the formation of these.

Sexual organs are not at present known in all the groups of
Ascomycetes. In certain orders they are entirely wanting, perhaps
as a result of reduction, so that the asci in these cases spring directly
from the mycelium.
2. The Basidiomycetes no longer possess sexual organs only in ;

the Uredineae or Rust-fungi are structures found which can be

regarded as persisting male organs, and cells which appear to
correspond to the carpogonia of Ascomycetes (cf. p. 406). Asci are
388 BOTANY PART n

not developed in Basidiomycetes, which are characterised by a special

kind of spore, the basidiospores. These are produced by a process of
budding on the characteristic BASIDIA, each of which bears a definite
number of spores, usually four. The typical basidia (Figs. 343, 344)
are either terminal tubular cells bearing the four spores at their
upper ends, or they are four-celled, each cell giving rise to a spore.
The spores are borne on thin stalk-like sterigmata, from which they
become separated when mature. The basidia agree with the asci in
containing when young two nuclei, which fuse with one another. The
7l
resulting nucleus then divides into the nuclei for the four spores ( ).
In most Basidiomycetes the basidia are borne on or in special
fructifications, the origin of which is not, however, dependent on a
sexual process.
The general name of CONIDIA is given to fungal spores which arise
by a process of budding, and the simple or branched hyphte bearing
them are termed conidiophores. The basidia are thus a special form
of the latter, but are distinguished from other conidiophores by their
peculiar nuclear fusion.
Other forms of conidiophore in addition to the basidia occur in
the life-history of some Basidiomycetes. In the Ascomycetes also
the formation of conidia as well as asci is a widespread method of
asexual reproduction. In general, it may be said that the asexual
formation of spores in the Fungi takes place in a large number of
different ways.
The direct origin of sporesfrom the cells of a hypha which round
off, become thick-walled and ultimately separate from one another
(chlamydospores of Brefeld), must be distinguished from the forma-
tion of conidia. The spores of the Ustilagineae are thus formed in
rows, while the uredo-spoues of the Uredineae arise at the ends of
hyphae.
All the asexual spores of the Eumycetes are provided with cell

walls, and are adapted for distribution through the air.

Sub-Class a*
I. Aseomyeetes (* .

TO)

The sexual organs have been accurately investigated in relatively

few forms a number of distinct types are found.
;

1. In the Laboulbeniaceae
(Fig. 342) the carpogonium with its
trichogyne, and the antheridia. which produce spermatia, show a
striking correspondence with the structures of the same name in
the Red Algae.
2. The Ascomycetes which enter into the composition of Lichens
(Figs.372, 373) approach most closely the preceding group. The
carpogonium is here a spirally wound filament of cells terminating in
'

a trichogyne. The spermatia are formed in special flask-shaped

depressions of the thallus, the spermogonia. Similar reproductive
SECT, i CRYPTOGAMS 389

organs occur in some Ascomycetes (e.g. Polystigma) which do not

form parts of Lichens.
3. Pyronema (Fig. 333) and related genera together with
Boudiera (
c2
)
exhibit a quite distinct type. A multinucleate carpo-
gonium which is provided with a trichogyne
is fertilised by a multi-

nucleate antheridium, the two structures being thus coenogametes.

4. In the Erysipheae (Fig. 328) a uninucleate antheridium unites

directly with a uninucleate oogonium.

It remains for future research to determine whether these types
are to be derived from a single one or whether they indicate a
polyphyletic origin for the Ascomycetes.
In many genera the sexual organs are more or less reduced, and
in some groups of Ascomycetes they are completely wanting.
Ascogenous filaments which bear the asci on their ultimate
ramifications (cf. Fig. 334) arise from the carpogonium. Sterile
hypha? which arise from below the carpogonium also take part in
the formation of the fructification, forming an investment to it and
paraphyses standing between the asci. The two sorts of hypha are,
however, clearly distinguishable.
The orders of Ascomycetes are provisionally characterised by the
construction of the fructification.
In the Perisporiaceae the small, spherical fruits (perithecia) are
surrounded by a complete investing layer, on the rupture or rotting
of which the spores are set free.
In the Discomycetes open cup-, club-, or hat-shaped fructifications
(apothecia) are formed the asci are arranged parallel to one another
;

termed the hymenium.

in a superficial layer
In the Pyrenomycetes the perithecia are flask-shaped, the asci
springing from the base of the cavity.
In the Tuberaceae the subterranean, tuberous fructifications are
closed.
To these orders must be added the Exoasci, in which the asci
arise from cells of the mycelium without the formation of any
and the very simple Saccharomycetes or Yeast-Fungi.
fructification,
These two groups can be regarded as extremely reduced Ascomycetes.
The Laboulbeniaceae in which the asci are enclosed in small peri-
thecia occupy an isolated position.

Order 1. Perisporiaceae (**<

(il
)

This order, which includes only Ascomycetes with enclosed fructifications, com-
prises two families the Erysipheae or Mildew Fungi and the Perisporieae.
:

1. The Erysipheae form a family of distinctive, epiphytic parasites whose

mycelium, somewhat resembling a cobweb, and ramifying in all directions over the
surface, particularly the leaves, of higher plants, sends out haustoria which
penetrate the epidermis of the host. In some cases the mycelium also inhabits
390 BOTANY PAUT II

the intercellular spaces of the leaf. The ripe ascus fructification! (perithecia) are
small black bodies provided with peculiar appendages. In the simplest forms
(e.g. in the genus Sphaerothcca) the spheroid perithecium encloses only a sin^lf
ascus with eight spores. It is enveloped by a covering of sterile hyphae, forming
a sheathing layer, two to three cells deep. The genera Eryripke and Uiicinula, on
the other hand, develop in each perithecium several asci, and in Plnjllactinia
12 to 25 asci are present. Since all the eight nuclei are not utilised in spore
formation the number of spores in each ascus is usually 4 or only 2. The peri-
thecia are irregularly ruptured at their apices and the spores are thus set free. As
HAI:I'KK lias shown, the first rudiment of the perithecium consists of an oogonium

tt ic-

Fio. 328. Sphaerotheca caxtaijnei. Fertilisation and development Of the peritheriuiii. 7, Ou-oninm

(o0) with the antheridial branch (02) applied to its surface S, separation of antheridium (an)
; ;

#, passage of the antheridial nucleus towards that of the oogonimn lt union of the nuclei
; , ;

5, fertilised oogonium surrounded by two layers of hypha; derived from the stalk-cell (st) ;6,
the multicellular ascogonium derived by division from the oogonium the terminal cell with
:

the two nuclei (us) gives rise to the ascus. (After HARPKR.)

and an antheridium. These are uninucleate cells, separated from the mycelium by
partition walls, and stand close together. The male nucleus passes into the
oogonium by an opening which forms in the cell walls (Fig. 328, 1-4). After
fertilisation the oogonium is surrounded by investing filaments which spring from
its stalk-cell or from that of the antheridium (5), and the oogonium itself becomes
converted into a multicellular structure (6). In Sphaerotheea the ascus containing

eight spores arises from the multinucleate terminal cell, while in Erysiphe and
Phyllactinia this cell produces ascogenous hyph; which in turn give rise to the
numerous asci.

Before entering upon the formation of perithecia, tin- .Mildew Fungi multiply
by means of conidia abstricted in chains from special, erect hyphae, from the tip
downwards. The Mildew Fungus of the Grape-vine occurs on the leaves and
berries in America and has appeared in Europe since 1845. This fungus, known as
Oidium Tuckeri, is the conidial form of Uiicinula necator (- U. spiralis], the small
perithecia of which have appendages spirally' rolled at their free ends and are only
rarely found (Fig. 329).
2. The Perisporieae are closely related to the Erysipln-ae, but are saprophytic
and live on decomposing organic matter. To this order belong two of the most
 CRYPTOGAMS 391

common Mould Fungi, Aspergillus (Eurotium), herbariorum and Penicillium

crustaceum. Both multiply extensively by means of conidia before they begin to
form perithecia.
In the case of Aspergillus herbariorum, the conidia are abstricted in chains from
a number of sterigmata arranged radially on the spherical, swollen ends of the
conidiophores (Fig. 330). The conidiophores are closely crowded together, and
constitute a white mould, afterwards turning to
a blue-green, frequently found on damp vege-
tables, fruit, bread, etc. Some species of Asper-
<l ill us are pathogenic in man and other mammals ;

thus A. fumigatus causes mycosis of the external

ear, the throat, and the lungs.
In Penicillium crustaceum, another wide-
spread blue-green mould, the erect conidiophores
(Fig. 330) are verticillately branched. The
spherical perithecia of Aspergillus and Penicil-
lium are produced later on the mycelium, but
only rarely occur in the latter genus. Their struc-
ture is more complicated than in the Erysipheae.
They originate as a spirally wound carpogonium

Fia. 329. Unciiiula necutoi: A, Conidial

stage ; c, coni<lium conidiophore. B,
; h,

Hypha which has formed ;i disc of attach-

ment (a) and has sent a haustoriuni (h) into
an epidermal cell. C, Perithecium with
appendages. (From SORAUER, LINDAU,
and REH. llani.lt>. cl. 2'flanzenl.rankheUfn, Fro. 330. Conidiophores of AsjiergUlus Jierlnirivrum (to the
ii. p. 194. lHOti.) left) and Penicillium cruxtaeeum (to the right.)

which after fertilisation becomes surrounded by lateral branches of the mycelium.

The ascogenotis hypha branches, penetrating the pseudo-parenchymatous tissue
within the perithecium, and bears numerous round asci containing eight spores.
The walls of the asci the surrounding pseudo-parenchyma disappear in the
ajid
ripe fructification which opens irregularly by the rupture of the peripheral layer.

48 ea
Order 2. Discomycetes (
-
)

The Discomycetes are distinguished from the other orders by their open
apothecia, which bear the hymenium, consisting of asci and paraphyses, freely
392 BOTANY

exposed on their upper surface (Fig. 332). The different groups exhibit great
diversity as regards the manner of development of their fructifications.

'Flo. 331. I'czlza aitrantiaca.

(After KKOMBHOI.Z, nat. size.)

Fio. 332. hichii'ii imlfherrima. Aixjthecium ruptured,

showing old and ynimx asri U-twecn tin- paiajihyses.
(After WOROXIN, from v. TAVEL.)

Fio. 333. Pyroiifma confluens. A, Rudiment of an apothecinm, cimsistin.; of three oogonia (og),
with tricho^ynes (t) and three antheridia (a). B, Fusion of the anttiei idium and the tip of the
trichogyne. (', The basal-wall of the trichogyne having been absorbed, the male and female
nuclei are ground in the centre of the ooKonium. D, Sejaration of the ootfonium from the
tricliox'.vne by means of a new jmrtition wall. Formation of the ascogenous filaments (as/). K,
Longitudinal section through a youn- ajujtlieciuin. u.; AMM. (After RHARPER. A, K
x about 150, B-D x about 300.)

The great majority of the Discomycetes, of which the genus Pezizti may serve
as a type, grow on living or dead
vegetable substances, especially upon decaying
 CRYPTOGAMS 393

wood, but sometimes also on humus soil. They produce saucer- or cup-shaped
fructifications of a fleshy or leathery consistency, and usually of small dimensions.
One of the largest forms, Peziza aurantiaea (Fig. 331), has irregularly bowl-shaped
fructifications, which be seven centimetres broad and of a bright orange-red
may
are grey or brown. Such cup-
colour, while in most of the other species they
sliaped fructifications are termed APOTHECIA.
The development of the apothecium may be described for Pyronema confluens,
in which it has been thoroughly investigated by R. HARPER. The fruit-body of
it often
this species is about 1 mm. across, and of a yellow or reddish colour ;

occurs on spots where fires have been kindled in woods. The carpogonia are
especially large in this species,
and several usually take part in the formation of
each apothecium (Fig. 333 A). The carpogonium or ascogonium consists of the

Kir;. 334. Development of the'Ascus. A-C,

Pyrone-iYM confluent. (After HARPER.)
D, Young ascus of Koudiera with eight
spores. ( After CLAUSSEN.) Explanation Fiu. 335. Morehella esculenta.
in text. (J nat. size.)

spherical, multinucleate oogonium, on the apex of which a multinucleate curved

cell, the trichogyne, is situated. The cylindrical, multinucleate antheridium
arises below the oogonium its apex comes into open communication with the tip
;

of the trichogyne by the breaking down of the intervening walls. The male nuclei
first wander into the trichogyne cell, and then, by the breaking down of the basal

wall of the latter, into the oogonium. In the oogonium the male nuclei conjugate
in pairs with the numerous female nuclei, while the nuclei of the trichogyne cell

disintegrate. The egg-cell then becomes limited from the trichogyne by a new
<(!! wall and sends out ascogenous filaments containing the conjugated nuclei.
These filaments branch and ultimately terminate in asci (E), while the sterile
hyphse and the paraphyses of the fructification are derived from hyphse arising
beneath the carpogonium. In Ascobolus, a genus related to Pyronema, the asco-
gonium is to begin with multicellular, but all the cells empty their contents into
a single large one, from which the ascogenous hyphse then arise.
The end of the ascogenous hypha ( 60 ) when about to form an ascus becomes
curved into a hook-like shape (Fig. 334 A). The two nuclei of the young ascus (a)
lie near to the bend, and on the formation of transverse walls are separated from
394 BOTANY PART II

the uninucleate terminal cell (/<) and the stalk-cell (s) which also has a single
nucleus. The two nuclei of the young ascus fuse ((7), and the resulting nucleus
gives rise by repeated division to the nuclei of the eight ascospores (Z)).
The highest development is exhibited by the peculiar fructifications of the
Helvellaceae, whose mycelium grows in the humus soil of woods. In the genus
Morchella (Fig. 335), the fructifications consist of a thick erect stalk, bearing a
club-shapeil or more or less spherical cap or pileus, which bears the hymenium,
with its eight-spored asci, on the reticulately indented exterior surface (Fig. 327).
The Morchellas are edible ( ta ), in particular M. csculenta and M. conica. The former
has a yellowish-brown cap, ovately spherical in shape, and attains a height of
12 cm. ;
is conical and dark brown, and it reaches a height
the cap of the latter
of 15 cm. Gyromitra esculenta, with dark brown cap and white stalk, and others
are also edible. In their external appearance the fructifications of these highly
developed Disromyretes greatly resemble those of the Basidiomycetes.

4 "-
Order 3. Pyrenomycetes (
4:
')

The I'yrenomycetes comprise an exceedingly varied group of Fungi, some of

which are parasitic upon different portions of plants, particularly on the cortex
and leaves, and others are saprophytic upon decaying wood, dung, etc., while a few
genera occur as parasites upon the larvae
of insects. The flask -shaped fructifica-
tions or perithecia are characteristic of this
order. The perithecia are open at the
top, and are covered inside, at the base,
with a Irymenial layer of asci and hair-like
paraphyses (Fig. 336). The lateral walls
are coated witli similar hyphal hairs, the
periphyses. The aseospoivs e>eape from
the perithecia through the aperture. In

Fio. 337. 7, Coniiliophcn- alisti ictinx conulin.

Fio. 336. Peritheciuni of l'odosporajlin't*f<lti from a pyenklinni of '
hypodenni'i.
c///''"*/""''
in longitudinal section. s, Asci ; a, (After BREFKI.D, x 300.) S, Pycnirliiim of>7riV-

paraphyses ; e, i>eriphysps ; m, mycelial g <>/,<lin;'ni in vertical section. (Aft>T

hyptiH-. (After v. TAVF.L, x 90.) TULASSK. x 70.)

this process one ascus after another elongates in consequence of the water absorbed.
and ejects its spores through the opening, or the sj>ores are set free within the
peritliecia. and are pressed out, embedded in a swollen mass of mucilage.
The simplest Pyrenomycetes possess free perithecia (Fig. 336) which are usually
 CRYPTOGAMS 395

small and of a dark colour, and grow singly on the inconspicuous mycelium (e.g.
if; Xplinfrin, and Podospord).
'

In other cases the perithecia are in groups.

H
Flo. 338. Clavieeps purjmrea. A, Mycelial hypha with conidia /.', car of Kye with several rip<-
;

sclerotia ; C, a sclerotium with stmmatu />. longitudinal section-of a fructification showing

:

numerous 'perithecia ; E, a single pi>ritlnciuiu, more highly magnifn-d F, ascus : with 'eight
filiform spores ; G, a ruptured ascus with vscaping ;S|H)IVS ; //. a siiij;lf siMirc. (A after
KREFELD ; C-H after TTI.ASNK : I! photograptie<l from nature. OFFICIAL ami I'oiso.vors.)

embedded in a cushion- or] club-shaped, sometimes branching, mass of compact

mycelial hyphae having a paeudo-parenchymatoua structure. Such a fructification
is known as a STROMA.
396 BOTANY PART II

In the life-history of most Pyrenotuycetes tin- formation of perithecia is

preceded by the production of various accessory fructifications, particularly of
-conidia, which arc abstricted in dim-rent ways, cither directly from the hypha>

-
c

Fir;. 339. Tuber nijinii. 1. A friictilicution in vertical section ( x 5); the cortex
,
:

d, air-containing tissue ; <. dark veins of compact liyplia- ; h, aseoxenous tissue :

3, a jiorticin ot'tlie hyineniiiin (x 460). (At't'-i TI I.ASXK. IVmn v. TAVEL.)

T

or from special conidiophores. and assist in disseminating the fungus. The

conidiopliores are frequently united in distinct, conidial fructifications. A
special form of such fructifications are the PVCNIIHA produced by many genera.
They are small spherical or Mask-shaped bodies which give rise to branched hyphal
filaments from the apices of which conidia, in this case termed PYCNOSPOIII.S or
are abstrictcd Fig. :$:}7, 1. 2).
, The structure of the pycnidia and
SECT, i CRYPTOGAMS 397

pycnospores corresponds to that of the spermogonia and spermatia of the Lichens,

and they may in part at least be regarded as male organs.

Claviceps purpurca, the fungus of Ergot, is important on account of its

official value. It is parasitic in the young ovaries of different members of the

Grarnineae, particularly of Rye. The ovaries are infected in early summer by

the ascospores. The mycelium soon begins to form conidia, which are abstricted
in small clusters from short lateral conidiophores (Fig. 338 A). At the same
time a sweet fluid is extruded. This so-called HONEY-DEW is eagerly sought by-
insects, and the conidia embedded in it are thus carried to the ovaries of other
plants. The conidial form of this Fungus was formerly regarded as a distinct
genus under the name of Sphacelia seyetum. After the completion of this form of
fructification, and the absorption of the tissue of the ovary by the mycelium, a
sclerotium is eventually formed in the place of the ovary from the hyphae of the
mycelium by their intimate union, especially at the periphery, into a compact
mass of pseudo-parenchyma (Fig. 107). These elongated dark-violet SCLEKOTIA,
which project in the form of slightly curved bodies from the ears of corn, are
known as Ergot (Fig. 338, B). The sclerotia, copiously supplied with reserve
material (fat), eventually fall to the ground, where they pass the winter, and
germinate in the following spring when the Rye is again in flower. They give rise
to bundles of hyphse which produce long, stalked, rose-coloured globular heads (C).
Over the surface of the latter are distributed numerous sunk perithecia (D, E).
Each perithecium contains a number of asci with eight long, filiform ascospores.
which are ejected and carried by the wind to the inflorescences of the grass.
Nectria ditissima C*4 ) is a very injurious parasitic fungus which inhabits the
cortex of various trees and causes the canker of fruit trees. It is propagated
in summer by means of slightly curved, fusiform, multicellular conidia, and
in winter and spring forms small red perithecia which are closely crowded

together.
OFFICIAL. Ergot is the sclerotium of ('/uric*'/,* jiti,-jni /''.

6S
Order 4. Tuberaceae (Truffles) ( )

The Tuberaceae or Truffle Fungi are saprophytic Ascomycetes whose mycelium

occurs in humus soil, particularly in woods. The ascus fructifications familiar
under the name of truffles are underground tuberous
bodies (Fig.. 339),
consisting
of a thick, investing layer, enclosing an inner mass in which the club-shaped
asci occur (Fig. 339, 2). The asei contain only a small number of spores in the ;

case of the true truffles (Tuber) they are usually only four in number, and

generally have a spinous or reticulately thickened epispore. When the fructifica-

tions are fully mature, the ripe spores lie free within the fructification.
3
The fructifications of many of the Tuberaceae C* ), and have an aromatic
are edible
odour and taste.
They are, for the most
part, obtained from France and Italy.
Of the edible varieties, the most important are the so-called black truffles belong-
ing to the genus Tuber, viz. Tuber brumale, mclanosporum, aestivuin, mescn-
tericum. The fructifications of these species have a warty cortex of a black,
reddish-brown, or dark brown colour. The white truffle, Choiromyces meanJri-
formis, the external surface of which is pale brown, is also edible.
Elaphomyccs granulatus (Boletus cervinus), the yellowish-brown fructifications
of which are of the size of a walnut and have a hitter taste, occurs commonly in
woods in Europe. It is used in veterinary medicine.
 398 BOTANY PART II

Order 5. Exoasci (**)

The most important genus of this group of Ascomycetes is Taphrina (including

Exoascus), the species of which are parasitic on various trees. They develop in
part annually beneath the cuticle of the leaves, causing discolorations of these
organs ; their mycelium persists during the winter in the tissue of the host, so
that a constant recurrence of the disease takes place. The presence of the
mycelium in the tissues of the infected part causes the abnormally profuse
development of branches known as WnriiKs'- BROOMS. Taphrina Carpini
produces the abnormal growths occurring on the Hornbeam Taphrina epiphylla,
;

those of .Units incana. Taphrina deformans

attacks the leaves of the Peach and causes them
-a*
to curl. Tii'/i/iriii-ii, Pruni is parasitic in the

young ovaries of many species of Prunus, and

produces the malformation of the fruit known
as "Bladder Plums," containing a cavity, the
so-called "pocket," in the place of the stone;
the mycelium persists through the winter in the
branches. In the formation of asci, which occurs
without the formation of sexual organs, in-
dividual cells of the copiously-branched mycelium
ramifying between the epidermis and cuticle
cut of the infected part become greatly swollen.
.

UOfc .en These grow into club-shaped tubes, which burst

through the cuticle and, after cutting off a
basal stalk-cell, are usually converted into asci
with eight spores (Fig. 340). The numerous
,,a I /(. Transverse asc i are
closely crowded together. In conse-
section through the epidermis of an .

infected plum, b our ripe asci, n <*., .

'
ce Q{ their increaaed t resulting from
}

with einht spores, 4 with yeast"

:! ,
an excessive absorption oi water, the asci become
like (-.onidia abstracted from the ruptured at their free extremities and eject the
st, stalk-cells of the asci
spores ;

m, filaments of the mycelium cut

transversely :
cut, cuticle; ep, epi-
;
spores.

.
^ L
-
whi(jh bu(j iu
wlution, frequently germinate while still en-
.
watw
...
Qr g
...

lemiis. (After SADEBECK, x 000.)

closed within the asci (Fig. 340, a 3 a 4 ), and ,

give rise to yeast-like conidia, e.g. in Taphrina Pruni.

by budding
The Exoasci are perhaps to be regarded as reduced Ascomycetes, in which the
sexual orj^iius have become completely suppressed.

67
Order 6. Saccharomycetes Yeast-Fungi) ^ ( )

The beer, and wine yeasts included in the genus Saccharomyces are
alcohol,
simple unicellular Fungi which assume the form of spherical, oval, or cylindrical
conidia containing a single nucleus. They increase in number by budding
(Fig. 341). No mycelium is formed, though sometimes the cells remain for a
time united in chains. With free access of oxygen and at a suitable temperature
yeasts form asci when the nutrient substratum is exhausted the asci externally ;

resemble the conidia but contain a few spores. In .some yeasts a conjugation of
two cells accompanied by a nuclear fusion has been observed ( 68 ). In Saccharo-
myces Ludwigii the four spores in the ascus germinate and fuse in pairs l>y means of
a narrow conjugation- tube the latter elongates into a germ-tube from which yeast-
;
 CRYPTOGAMS 399

cells are abstricted. In the ginger -beer yeast (Zyyosacckuroinyccs} and in

Schizosaccharomyces the yeast cells conjugate by means of long tubes before
-

spore formation.
- These nuclear fusions possibly correspond to those in the
young asci of other Ascomycetes.
Physiologically these Fungi are remarkable for their power of exciting, by
means of an enzyme (zymase), the
fermentation of saccharine solutions,
alcohol and carbon dioxide being
produced. The beer yeast (tiacchci-
romyces cerevisiae) is only known in
the cultivated form the wine yeast
;

(S. cllipsoideus), on the other hand,

occurs regularly in the soil of vine- p
A, yeast-cell ; B, C,
yards in the spore-form the latter
;
yeast - cell budding ; D, ascus with four spores.
is therefore always present on the
(After GUILLIEUMOND, x 1125.)
grapes and need not be added to the
grape-juice. Other genera, in some of which a mycelium is developed, belong to
this order.
No evidence is at present forthcoming to show that the Yeasts are to be
regarded
as developmental forms of other

Fungi. In various members of the

Exoasci, and Ustilagineae, however,
yeast-like conidia which reproduce
by budding are known. Possibly
the Saccharomycetes are reduced
Ascomycetes, or they may represent
an independent group of very simple
Fungi.

Order ro
7. Laboulbeniaceae ( )

Tlu; Laboulbeniaceae are a group

of minute Fungi occupying an iso-
lated position our knowledge of
;

them is
largely due to the work
in-
of TH AXTEU. Their thallus consists
of two to a number of cells, and
is attached to the body of the in-
sect, most commonly a beetle, on
which it is parasitic by means of
a pointed process of the lowest cell
inserted into the chitinous integu-
ment of the insect or by means of
rhizoids which penetrate moredeeply.
Fie. S&.-StignMtomyccs Jiaeri. Description in text. Utiyniatomijces Baeri which occurs
.4, spore;B-F, sucees>i\v developmental stages. D. onhouse-fliesin Europe may be taken
with spermatia escaping from the antheridia are ; K, as an
example. The bicellular spore
with antheridia above and the lateral female organ ;
F, Perithecium with developing asei <1, ripe ascus.
;
(pig> 342 ^
which has ft mucilagi .
nous outer coat, becomes attached
(After THAXTER.)
by its lower end (B), and divisions
occur in both [cells (C). From the upper cell an appendage is developed bearing
400 BOTANY

a number of unicellular, flask-shaped antheridia (D, an) from \\liii-li naked

spherical spermatia without cilia are shed. The lower cell divides into four
(D, a, b, c, d), and the cell a projects and gives rise to the multicellular female
organ. The true egg-cell (E, ac), which is called the carpogoiiium. is surrounded
by a layer of cells. Above the carpogonium come two cells (E, tp, t) the upper of
which is the freely projecting trichogyne or receptive organ for the spennatia.
The behaviour of the nuclei has not yet been followed, but probably a fusion of
male and female nuclei takes place in the carpogonium. The latter becomes
divided into three cells of which the uppermost disappears, the lowest (F, st)
remains sterile, while from the middle cell the asci grow out. Each ascus ((/)
produces four spindle-shaped, bicellular spores. The similarity of the sexual
organs to those of the Florideae and of the lichen-forming Ascomycetes is of great
interest.

48. 49. 70. 71- 72

Sub-Class II. Basidiomycetes (

The large group of the Basidiomycetes in the wide sense consists

of Fungi, the mycelium of which is septate as in the Ascomycetes, but

Flo. 343. AriiiiHin-iii uniltti. .1, Yonii.^

Uisidiuin with the two primary
nuclei ; B, after fusion of the two
nuclei. Hypholomn /'/" '"'" "la-turn,

C, a liasidinm before the four nuclei Kl(i. :i44.--Uasi.lin. .1. Of a rrcdineoiis

derived from the secondary nucleus fungus (EiKlii/ihiillinti I-'.HI i/i M ;'.'

of the basidium have passed into the .

i.aftei-Tri.AsxE). /:>', Of one of the
four iKisidiospores 1). passage of a
; Treniellineae (Trrmelln lnti'*>; HX) (after
nucleus through the sterigma into HKEKKI.HX (x 450. From v. TAVEI,,
the liasidiospore. (After RUHLAXD.)

is distinguished by the loss of sexual reproduction and by the

formation of basidia (p. 388), different forms of which are met with.
In the orders Uredineae and Auricularieae the upper portion of
the basidium is divided by transverse walls into four cells ; each cell
bears on a thin stalk (sterigma), arising near the upper end, a single
spore (Fig. 344 A}. In the Treniellineae, on the other hand, the
basidium is divided by longitudinal walls into four cells, each of
which continues into a long tubular sterigma (Fig. 344 B). The
 401

basiclium in the Hymenomycetes and Gasteromycetes is unicellular,

and bears as a rule four spores at the summit these may be sessile ;

or situated on sterigmata (Figs. 343, 355).

The Ustilagineae are of interest since in one family the basidia
are divided, and not always strictly four-celled, while in the other
the basidia are undivided. The number of spores produced is not a
definite onebut often very large.
In addition to conidiophores differentiated as basidia, the Basidio-
mycetes produce other forms of conidia as accessory fructifications in
many species. These appear in the Ustilagineae as the smut-spores,
and as the rust-spores in the Uredineae. In these groups the basidia
arise directly from spores of this kind (Figs. 344 A, 346). In other
Basidiomycetes, if a few simple forms are disregarded, the basidia are
always borne upon or within more or less complicated fructifications.
The layer in which the basidia are associated together is termed the
hymenium.
48 49 . 73
Order 1. Ustilagineae (Smut-Fungi) (
-
)

The Ustilagineae are parasites, and their mycelium is found ramifying in higher

plants, usually in definite organs, either in the leaves and stems, or in the fruit or
stamens. The Gramineae in
particular serve as host plants ;

certain species of Ustilagineae

are in a high degree injurious
to cereals, and produce in the
inflorescences of Oats, Barley,
Wheat, Millet, and Maize the
disease known asSmut.
The mycelium ultimately
produces resting-spores by the
formation of additional trans-
verse walls, and by the divi-
sion of its profusely branched

hyphse into short swollen cells

(Fig. 345 A). The cells be-
comeroundedoffand converted
FIG. 345. .!, Ustilago olivacea. Mycelial hypha in process of
into spores, within a gelatinous
forming smut-spores ( x 400). B-D, Ustilago segetum: B,
envelope, which, however, germinating smut-sport- (i-l), cultivated in nutrient solu-
The tion t, transversely septate, liasidiuin with lateral and
eventually disappears. ;

terminal basidiospores (conidia) (e) ( x 450); C, germinating

spores then become invested smut -spore lying in the nutrient solution surrounded
with a new, thick, double
by ab.stricted conidia, which are multiplying by budding
wall. In this way the mycel- ( X 200) I), an aggregation of budding conidia ( x 350).
;

ium is transformed into a dark (After BREFELD, from v. TAVEL, Pilze.)

brown or black mass of spores.
These Smut-spores, brand-spores, or resting-spores are scattered by the wind, and
germinate only after an interval of rest, producing the basidia in the succeeding
spring the formation of these is characteristically different in the two families of
;

the Ustilaginaceae and the Tilletiaceae.

The most important genus of the Ustilaginaceae is Ustilago. Ust. Avenae,
2 D
402 HOT A X Y

U. Hordei, and I'. which were formerly united as U. Carbo, cause

Tritici sf yet urn,
the "smut" or of Oats, Barley, and Wheat.
"brand" The mycelium penetrate>
the ovary, and forms dark brown, dust -like masses of escaping resting -spores.
U. Afaydis produces on the stalks, leaves, and inflorescences of the Maize tumour-
like swellings filled with brand-spores in the form of a black powder. Other >peei.-s
live on the leaves of different grasses while U. violacea ( = U. antherarum) occurs
;

in the anthers of various Carophyllaceae (e.<j. /,//<///., Saponaria), and fills the

pollen-sacs with brand-spores.

The brand-spores of Ustilago fall to the ground, and after a period of rest give
rise, on germinating, to a
short tube (promycelium) which becomes septatcd \>\
three or four transverse walls (Fig. 346 ), and,
functioning as a basidium, produces ovate basidio-
spores or conidia (sporidia), both laterally from the
upper ends of the intermediate cells, and also from
the tip of the terminal cell. When abundantly
supplied with nourishment, as when cultivated in
a nutrient solution, conidia are
continuously ab-
stricted in large numbers (Fig. 345 (7), and then
multiply further by budding, cell-fusions or union
of nuclei frequently occurring as a preliminary (C,
D}. After the food-supply of the substratum is
exhausted, the conidia grow out into mycelial hyphse.
The formation of the conidia in the damp manured
soil of the grain fields is accomplished during a
saprophytic mode of existence, but the hyphal fila-
B ments which are eventually produced become para-
FIG. 34i; sitic an d penetrate the young seedlings as far as the
.
i'xtilugo Scabosine. A, >

yoiiiij; hasidiuni with four nuclei apical cone where the inflorescence takes its origin,
formed on germination of the T ue mycelium continues its development in the
resting spore S, spore-formation
;

inflorescence) and ultimately terminates its exist-

(in the 4-ei'lh-il hasiilium. (After
HARPER.)
ence by the production of brand-spores. No conidia
are formed on the host plant itself.
In addition to the infection of young plants either resting-spores or the conidia
4
resulting from their germination may be carried to the stigmas of the grass- flowers (~ )
and germinating there produce a mycelium which penetrates to the young seed>
and passes the winter in the embryo-plants. Such infection of the flowers may
alone take place as in Ustilago Tritici, U. Hordei, and U. antherarum, or the seed-
ling may more often be infected as in U. Avenae, U. Sorghi, U. Panici miliacei,
U. Crameri. The Smut of Maize can infect all parts of the plant while in a young
state and the disease is limited to the
infected spots.
The life-history of the Tilletiaceae
is similar to that of the Ustilaginaceae. The
best-known species are Tilletia Tritici (=T. Caries] and Tilletia laevis, the fungi
of the stink-brand of wheat. The resting-spores till the apparently healthy grains
and smell like
fish. In the first- named species the resting-spores are
decayed
reticulately thickened those of T. lacvis, on the other hand, are smooth-walled.
;

Unlike the Ustilaginaceae, the germ-tube gives rise only at its apex to filiform
basidiospores, which are disposed in a whorl, and consist of four to twelve spores
(Fig. 347 A). The basidiospores also exhibit the peculiarity that they coalesce
with one another in pairs in an H-form. Such cell fusions also occur between
germinating conidia of the Ustilagineae, but are not accompanied by nuclear fusions.
The filiform spores germinate readily, and produce sickle-shaped conidia at the apex
 CRYPTOGAMS 403

of the germ- tubes (Fig. 347 C). When abundantly supplied with food material, the
germ-tubes grow into large mycelia, from which such sickle-shaped conidia are so
abundantly abstricted that they have the appearance of a growth of mould (Z>).

FIG. 347. Tilletia Tritici. A, the basidium developed from the brand-spore bearing at the end
four pairs of spores A: (x 300) B, the dispersion of the spores which have fused in pairs ( x
;

250) C, one of the paired spores germinating and bearing a sickle-shaped conidium sk (x 400).
;

D, Mycelium with sickle-shaped conidia (x 350). (After BREFELD.)

Thus Tilletiii, unlike produces conidia of two forms

Ustilago, but in other ;

particulars the development of both groups is the same.

The young resting-spores, and the cells of the mycelium from which they are
produced contain each two nuclei which fuse with one another as the spore becomes
mature. The cells of the basidia, and the basidiospores are all uuinucleate, only
the secondary conidia of Tilletia being again binucleate.

Order 2. Uredineae (Rust Fungi) p-?8 )

. The mycelium of the Uredineae lives parasitically in the intercellular spaces of

the tissues of the higher plants, especially in the leaves, and gives rise to the
widely spread diseases known as Rusts. Their more varied spore- formation is a
distinguishing feature as contrasted with the Ustilagineae.
As in the latter order, the basidia are not produced directly on the mycelium
but on the germination of a special type of spore, teleutospores or winter spores,
which are characteristic of all Uredineae. The teleutospores arise in small clusters
beneath the epidermis of the diseased leaf from the ends of hyphse frequently ;

two or more form a short chain. They are thick- walled resting-spores and persist
through the winter (Fig. 348, 1, ot). The group of spores usually bursts through
the epidermis. At first the spores, like the cells of the mycelium which bears
them, have two nuclei, but the nuclei fuse before the spore is ripe.
In the germination of the teleutospore a basidium (promycelium) grows from
each cell (Fig. 348, 2) it becomes divided by transverse septa into a row of four
;

cells from each of which a sterigma bearing a single uninucleated basidiosjwre

(sporidium) is produced. The sporidia are dispersed by the wind and germinate
in the spring on the leaves of host plants (which may be of the same or different

species from the one on which the teleutospores were produced), giving rise to an
intercellular mycelium, all the cells of which are uninucleate. From this
mycelium organs of two kinds arise, spermogonia on the upper surface of tin- leaf
and secidia on the lower surface.
2 D 1
404 BOTANY

The SPEEMOGOKIA (Fig. 349) are flask-shaped structures, the base of which is

covered with the projecting ends of hyphse from these are abstricted spermatic.
;

each of which has a single nucleus. Morphologically they arc completely i- <

parahle to the similarly named male sexual organs of some Ascomycetes amon^ ;

the Basidiomycetes they persist only in the Uredineae, and even in them are no

FIG. 348. Puccinia graminis. 1, Transverse section through;^ grass-haulm with group of teleiito-

spores ; germinating teleutospore with two basidia

?, ; vegetative. //, fructifying gciminatin;:
.-',

basidiospore tin-' latt'T has formed a secondary spore,, not having been able to infect a lio.-t
;

plant ./, a ]KH tion of a group of uredospores ()

; and teleutosporcs (() p, the germ-spores
; : '.'.

germinating uredospore. (1, 2, 3, U, after TUI.ASXE ; 6, after DK BARY

.'>, ; 1 x 150, 2 x circa

230, 3, 4 x 370 5 x 800, 6 x 390, from v. TAVEI., /'//:..)

longer functional and may be completely wanting. In nutrient solutions the

spermatia may put out short germ-tubes but are not capable of infecting the h"st
plant.
The JECIDI\ (Fig. 350) are cup-shaped fructifications, which when young are
closed, but later open from the ends of the hypha- numerous closely associated
;

chains of spores are abstricted. As a rule the enveloping layer or pcridium of the
 CRYPTOGAMS 405

lecidium is formed of thick-walled cells corresponding 'to the sterilised peripheral

rows of spores. In Phragmidiiim violaceum, which occurs on Blackberry leaves
and has been fully investigated
by BLACKMAN
76
( ), the hyphen
beneath the epidermis when
about to give rise to an secidiuni
cut off a sterile cell, which
iir.st

undergoes no further develop-

ment, from their ends (Fig. 351
A}. The cell below this increases
in size it has at first only a
;

single nucleus but becomes bi-

nucleate by the passage of a
nucleus into it from an adjoining
mycelial cell. The two nuclei do
not fuse. The bimicleate cell

undergoes successive divisions

into achainof spore-mother-cells,
each of which has a pair of nuclei;
FIG. 349. Gymnoxporungium davariacforme.
and from each spore-mdther-cell goniuni rupturing the epidermis of a leaf of Crataegus ;

an upper binucleate aecidiospore sp, spennatia p, sterile paraphyses. (After BLACKMAN.)

;

and a sterile intercalary cell,

which is also binucleate but soon shrivels up, are derived by a transverse
division (B, O).

Fio. 350. I'uccinia graminis. /Ecidium on Herberts vulgaris ; </, epidermis of lower surface of leaf ;

I'm, intercellularlmycelium yep, peridiumi; ,

chains of spores. (x!142.)

76 the development of the secidiospores in Phrag-

According! to ^CHRISTMAN ( )

midium spetiosum (Fig. 352), which is parasitic on Rosa, proceeds somewhat

differently. Here also the ends of the hyphse (A) divide into a terminal sterile cell
406 HOT A N V I'AIIT II

and a lower fertile cell (/>'), hut the fertile cells fuse in pairs with one another.
the upper i>ortions of tin- separating walls breaking down (C). The two nuclei lie
side by side and divide simultaneously (conjugate division. Two of the daughtei-
miclei remain in the lower part and two pass to the upper portion of the dividing
cell, and this upper portion is
separated hy a transverse wall as the first spoiv-
mother-cell (/>). In other respects the formation of the secidiospores proceeds as
described above. A iwridium is not formed in Plirmjiiiiilni.m.
The ripe, hinucleate ;ecidiospores (Fig. 3f>l
D) arc shed and infect a new host plant. Kadi
spore gives rise to an intercellular mycelium
which soon proceeds in the summer to hear uredo-
spores or summer spores. These appear in small
circular or linear groups and arise singly from
the enlarging terminal cells of the hyplue (Fig.
348, 5, 6). They have two nuclei like all the cell-
of themycelium developed from the secidiospore.
They commonly to ensure the spread of the
serve
fungus in the summer. Later, either in the same
or in distinct sori the teleutospores are formed.
and in these the fusion of the two nuclei to a
single one takes place such a fusion as a rule
;
is

found to take place in the young basidium.

The two types of cell-fusion in the formation
of the recidium are also found in other Uredineae,
and must be regarded as replacing a formerly
existing method of fertilisation. If we attempt
to derive the Uredineae from the Ascomycetes
the spermatia must be regarded as now function-
less male cells, and the so-called fertile cells in
the young secidium as corresponding to carpo-
gonia. Extending the comparison further the
mycelium proceeding from the fecidiospore in the
FIG. 351. Phrngmiilium riolaeeum. Uredineae and the nredospores and teleutospores
portion of a young tecidium , ., ... ,, ,. , . ,.

sterile cell fertile cells at <* bonle n lt to gether wlth the basldia formed
>
>
; , ;

tli.- ]>;.ssage of a nucleus from the by the would together correspond to the
latter,
adjoining cell is seen ; B, formation asexual of the Asco-
generation (sporophyte)
of the first spore-mother-cell sm,
niyce tes The basidiospores would thus correspond

which from i the from the basidiospore would be equivalent to the

first iecidios^re
() and the intercalary cell (z) have
sexual generation (gametophyte).
arisen ; smo, the second spore-mother-
flie life-history of the Rust-fungi is thus a
WCidiOSI> rP (Am>F
complicated one. The several
'
forms of spore may
B^cRMA^r
of the year on the one host.
appear in the course
such Uredineae termed autxecious. On the other hand the spermogonia and
being
jecidia occur on one species of host plant, and the uredospores and teleutosiKjrcs
may
on another, often unrelated, plant. In these hetercecious species there is thus an
alternation of host plants ( 77 ).
An example of an hetercecious Rust-Fungus is afforded by Pweiltia pnMRMlM,
the rust of wheat. It develops its uredospores and teleutospores
on all the green
of Gramineae, of Wheat, Barley and Oats. The a>cidia and
parts especially Rye,
of the Barberry
spermogonia of this species are found on the leaves
 CRYPTOGAMS 407

vulgans). In the spring the hibernating double teleutospores give rise to

transversely septate basidia, from which the four basidiospores are abstricted
(Fig. 348, 2). These are scattered by the wind, and if they fall on the leaves of
the Barberry they germinate at once. The germ-tube penetrates the cuticle, and
there forms a mycelium which gives rise to spermogonia on the upper side of the
leaf,and to fecidia on the under side (Fig. 350). This form of the fungus is
known as Ai'dilium Berberidis. On the rupture of the peridium the reddish-
yellow fecidiospores are conveyed by the wind to the haulms and leaves of grasses,
upon which alone they can germinate. The mycelium thus developed produces
at first uredospores (Fig. 348, 5). They are unicellular, studded with warty
protuberances, and provided with four equatorially disposed germ-pores. In
of the reddish-yellow fat globules contained in the protoplasm of

C
FII;. :;">-. A, The first rudiment of an a-cidimn beneath the epidermis of
I'liriiiiiitliliniii *iir,-i<>xi<ni.

a leaf of Rosa R, the division of the end-cell of a hypha into the upper, transitory, sterile cell
;

and the lower fertile cell C, conjugation of two adjoining fertile cells D, later stage in which
; ;

the first nuclear division is completed E, abstriction of the first fecidiospore mother-cell F,
; ;

chain of svcidiospores (a^ 2 ) separated by intercalary cells (Z], z) sm, the last-formed spore-
;

MKither-cell still undivided. (After CHRISTMAS.)

the spores, the fructification forms red markings (formerly termed Uredo lincaris)
cm the epidermis of the host plant. The uredospores are capable of germinating
at once on the wheat, and thus the rust disease is quickly spread. Towards the
end of the summer the same mycelium produces the dark brown, thick-walled
teleutospores, which in this species are always double, being united in pairs.
Each teleutospore is provided with one germ-pore, and on germination in the
succeeding year the cycle is begun afresh. The mycelium of the Uredo form may
hibernate in winter wheat, and thus the rust may appear in the spring without
the previous formation of basidiospores or of aecidia.
Other common Rusts on grasses or cereals which are closely related to Puccinia
graminis are Puccinia dispersa on the Rye, which develops its secidia on Anchusa,
and P. triticina and P. <jhi miirvii), which occur respectively on. Wheat and on
\Vlicat, Rye and Barley. The a?cidia of these latter forms are unknown.
All Uredineae do not exhibit so complicated a course of development as Puccinia
iirn nt iii is.Certain species produce only basidia from germinating teleutospores,
'or, in addition to teleutospores, only uredospores which are developed on the same
2 D 2
408 BOTANY

host. Others produce spermogonia and aecidia, and afterwards teleutospores, but
no uredospores. In the case of the hetercecious species, it is only possible to
demonstrate the connection between the different spore-forms by means of culture
experiments. So long as the relation of the different forms was not known, it was
customary to designate each by a special generic name the Uredo forms as Uredo ; ;

the jEcidia, according to their structure, as Aecidium, Boestelia, Peridermium ,

etc. The generic name is now determined by the characters of the teleutospores,
since theyexhibit the most characteristic distinctions.
In those Uredineae which no longer possess secidia and spermogonia, the cells of
the vegetative mycelium arising from the basidiospore are uninucleate, but subse-
quently, before the formation of the teleutospores. binucleate cells are found.
Further research is needed to show how the binucleate condition is arrived at ;

at all events, these forms exhibit a still further reduction of the sexual process and
of the alternation of generations.

Order 3. Auricularieae

The basidia, as in the case of the Uredineae, are transversely septate, with four
spores. Only a few forms are included in this order. Among the most familiar is
Auricularia sambucina (Judas' ear), found on old Elder stems. It lias gelatinous,

dark brown fructifications, which are shell-shaped and bear on their inner sides
the basidial hymenium.

Order 4. Tremellineae

The basidia are longitudinally divided (Fig. 344 JJ). The hymenium is
situated on the upper surface of the fructifications, which are generally gelatinous
and irregularly lobed or folded.
The few genera included in
this order are saprophytic on

decaying wood and tree-

trunks, on the surface of
which the fructifications are
produced.

Order 5. Hymenomy-
Kl 7
cetes (
-

'-)

The basidia are undivided,

and bear four spores at the
apices of slender sterigmata
(Fig. 353 sp). In the simplest
forms these basidia spring
directly from the mycelium,
but in the great majority of
FIG. 353. Russula rubiii. Portion of the liynifniuin. xh, Sub- cases fructifications are pro-
hymenial- layer ; ?/. liasidia ., sterigmata
; ; *r. s]mn-s ;
/',
which bear definite
duced,
paraphysps c, a cystid. (x 540.)
;

hymenial layers, comjiosed, in

addition to the basidia, of paraphyses (Fig. 353 p), and also of sterile cystids (c)
or club-shaped tubes characterised by their larger diameter and more strongly
thickened wall.
Most of the Hymenomycetes develop their profusely branched, white mycelium
SECT. I CRYPTOGAMS 409

in the humus soil of forests, in decaying wood or on dying tree trunks, and
produce fructifications, commonly known as toad -stools, protruding from the
substratum. The mycelium of the forms vegetating in the soil spreads farther
and farther, and dying in the centre as it exhausts the food material of the
substratum, occupies continually widening concentric zones. In consequence of
this mode of growth, where the development 'has been undisturbed, the fructifi-
cations,which appear in autumn, form the so-called fairy rings. A few Hymeno-
mycetes are parasitic, and vegetate in the bark or wood of trees. Of such parasitic
forms Armillaria mellea, whose mycelium vegetates between the bark and wood of
Conifers and other trees, is a familiar example. The profusely branching mycelial
hyphae (Fig. 354) become interwoven into flat, black strands from which tine, hair-

Fio. 354. -\nnilliii-iii mi'Ui'u. Por-

tion of a rhizomorph strand (>)
with mature (n) and youn<; (')
fructifications. (After HARTIO,
from v. TAVEL ; \ nut. si/.-.)

like hyphte are sent out and penetrate the wood for the absorption of nourishment.
It is from these sub-cortical strands, known as IIHIZOMORPHA, that the stalked,

capitate fructifications are eventually produced. In addition to the subcortical

strands, subterranean rhizomorphs are developed which pervade the soil and
infect the roots of other trees. The rhizomorphs may l>c regarded as a form of
sclerotia. This fungus is one of those that give rise to photogenic substances
which cause the phosphorescence of the infected wood ( 79 ).
The Hymenomycetes are further classified according to the increasing complexity
exhibited in the structure of their basidial fructifications.
1. In a few genera no distinctive fructifications are formed, and the basidia

spring in irregular groups directly from the mycelium. Exobasidium l"accinn may
be taken as a type of this form. The mycelium of this fungus, which is widely
spread in Europe, is parasitic in the Ericaceae, especially on species of Vaccinium
;

itcauses hypertrophy of the infected parts. The basidia are formed in groups
under the epidermis, which they finally rupture (Fig. :j ;>"). In this genus, as in
410 BOTANY PAKT 11

many others, accessory fructifications are developed, ami spindle-shaped conidia

are abstricted from the mycelium on the surface of the host plant, before tin
formation of the basidia.
2. In the group of the Thelephoreae. distinctive fructifications of a simple type
are found. Tln-y form on the trunks of trees either flat, leathery incrustations
liearing the hymenium on their
smoot upper surfaces; or the flat
li

fructifications becomeraised above

the substratum and form bracket-
winch frequently
like projection*,
show an imbricated arrangement,
and bear the hymenium on the
under side(e.g.Stcrcumhirsitti/n/,
common on the stems of decidu-
ous trees).
3. The fructifications of the

Clavarieae form erect whitish or

yellow
- coloured bodies, either

fleshy and club-shaped or more or

le>s branched, in a coral-like
fashion. The
larger, profusely
ln-anehed forms of this group are
FIG. 3.j(j. i'lararia botrytis. (X;tt. si/t>.)
highly esteemed for their edible
qualities in particular. Clavaria flava, whose fleshy, yellow-coloured fructifications
;

are often ten centimetres high, also Clavaria botrytis (Fig. 356), which has a pale
has fructi-
red colour. Sparassis crispa, which grows in sandy soil in Pine woods,
fications half a metre in diameter, with compressed, leaf-like branehes.
4. The Hydneae have fructifications with spinous projections over which the
hymenium extends. In the simpler forms
the fructifications have the appearance of
incrustations, with spinous outgrowths
projecting from the upper surface in other ;

cases they have a stalk, bearing an umbrella-

like expansion, from the under side of
which the outgrowths depend. The latter
form is exhibited by the edible fungi
Hydnum imbricatum, which has a brown
pileus 15 cm. wide, with dark scales on the
upper surface, and Hydnum repandum(Yig.
357),with a yellowish pileus.
5. In the Polyporeae, a group containing
numerous species, the stalked or sessile and
bracket-shaped fructifications are indented
on the under side with pit-like depressions, ,,'iinliiiii.

or deep winding passages, or covered with

a layer of tubes, closely fitted together and lined by the
hymenium. To this family
belongs the genus Boletus, occurring on the soil of woods, which has a large, thiek-
stalked pileus, covered on the under side with a layer of narrow dependent tubes.
Although many species of this genus are edible (e.g. B. edulis), others are
exe.-edingly poisonous, in particular Jj. Satanns (Fig. 360). The stalk of the latter
fiingus is a yellow to reddish-purple, or has red reticulate markings, while the
 CRYPTOGAMS 411

pili'iis, which may bo 20 cm. wide, is yellowish-brcnvn on its upper surface, but on
tin- iiii<lt'v side is at first blood-red, becoming later orange-red. Of the numerous
spi-cics of the genus Polyporus, P. fomcntarius, Touch-wood, is used in medicine.

Km. :r.8.--7W///i/-.< tffniaritu. Si-ction through an old Fio. 359. Pmlliota eamjientria ( = Aga-
frustification, showing annual /ones of j^'owth a, ; rlciis campestris). To the right, a
liciint of attachment. (J nat. .size.) yonnj; t'l uctiliraticm. (Reduced.)

Its mycelium is parasitic in deciduous trees,

especially the Beech, and produces
large, bracket or hoof-shaped, perennial fructifications, 30 cm. wide and 15 cm.
thick. They have a hard, grey external surface, but inside are of composed softer,

Km. 3i>0. * Sutunrtx. (After KROMBHOLZ, A nat. size.) 1'OISOXOCS.

more loosely woven hyphse, and were formerly used for .tinder. The narrow tubes
of the hymenium are disposed on the under side of the fructifications in successive
annual layers. /'.
iij/t.-iarivs (Fig. 358), which is often found on Willows, and has
a similar structure, has a rusty-brown colour, and furnishes, since it is much harder,
a poorer quality of tinder. Polyporus officinalix with an irregularly tuberous
412 BOTANY

white fructification occurs on Larches in South Europe ;

it contains a liitti-r

resinous suli-tancc amiis also used in medicine.

Many parasitic 1'olyporeae are highly injurious to the trees attacked by them ;

thus Jfcterobasidion annosum often cause* the death of whole forests of 1'iiies and
Spruce Kirs. .)/< nilins
lacrymans C*) is an exceedingly dangerous saprophytio
attacking and destroying the timber of damp houses.
s}>ecies, The niyeeliuni of
this fungus forms large, wliite, felted masses with firmer branched strands which
serve to conduct water and food substances. It gives rise to outspread, irregularly

shaped, pitted fructifications of an ochre or rusty-brown colour, and covered with a

hymenial layer. Good ventilation
of the infected space is the best
remedial measure.
6. The Agaricineae, which in-
clude the greatest number of species,
have stalked fructifications, com-
monly known as Mushrooms and
Toadstools. The under side of the
pileus bears a number of radially
disposed lamelhe or gills which are
covered with the basidia- producing
hymenium. In the early stages of
their formation the fructifications
consist of nearly spherical masses
of interwoven hyphre, in which the
stalk and pileus soon become differ-
entiated. The rudiments of the
stalk and pileus are at first enclosed
in a loosely woven envelope, the
VOLVA. In the course of the further
development and elongation of the
stalk the volva is ruptured, and its
torn remnants form a ring or sheath
at the base of the stalk, and in many
the "Fly Mushroom,"
cases (as in
FIG. 361. Amanita muscaria. (Jnat. size.) I'oisoxvi's. Amanita muscaria), are still trace-
alile in the white scales conspicuous

In addition to the volva many Agaricineae

on the red surface of the pileus (Fig.361).
which
develop a so-called VELUM, consisting of a thin membrane of hyphal tissue
extends in young fructifications from the stalk to the margin of the pileus, but is
afterwards ruptured, and remains as a ring of tissue encircling the stalk (Fig. 359).
Many Mushrooms found growing in the woods and fields are highly
of the
esteemed as articles of food. Of edible species the following may be named: the.
common Field-Mushroom, now extensively cultivated, Psalliotn a, mjnxtris (Fig.
359), with whitish pileus and lamellae at first white, then turning flesh-colour,
and
finally becoming chocolate-coloured ; Cantharellus cibarius, having
an orange-
coloured pileus Lactarius dcliciosiis, which has a reddish-yellow pileus, and con-
;

tains a similarly coloured milky juice in special hyphal tubes Lcpiota procera, ;

whose white pileus is flecked with brown scales Amanita cacsarea with an orange
;

pileus bearing a few white scalesand yellow lamellae.

Of the poisonous Agaricineae the following are best known Amanita nuts*' rid :

(Fig. 361); Amanita bulbosa (Fig. 362) with whitish or yellowish pileus
and the
 CRYPTOGAMS 413

stalk swollen at the base Russula emctica, with a red pileus and white lamellae
; ;

Lactarius torminosus, having a shaggy, yellow or reddish-brown pileus and white

milky juice.
Rozites yongylophora,
found in South Brazil, is of
especial biological interest.
According to A. HOLLER, this
species is regularly cultivated
in the nests o f the leaf-cutting
ants. Its mycelium produces
spherical swellings at the ends
of the hyphse, which become
filled with protoplasm, the
so-called Kohl-rabi heads,and
serve the ants as food material .

The ants prevent the develop-

ment of the accessory conidial
fructifications peculiar to this

fungus, and thus continually

maintain the mycelium in
their nests in its vegetative
condition. The fructifica-
tions, which rarely occur on
the nests, resemble those of
Amanita muscaria, with
which Rozites is nearlyallied.
AccordingtoHoLTERMAN,the
mycelium of Agaricus rajap
is cultivated in their nests by

termites in tropical Asia ( 81 ).

ECONOMIC USES.- Poly-
FIG. 3C2. Aiiianita Imlbosa. (J nat. size.) Potsoyoi'f.
porus fomentarius (FUNGUS
CHIRURGORUM). Polyporus
officinalis (
Boletus laricis) gives AGARICUS ALBUS, AGARICINUM, and ACIDUM
AGARICINUM.

Order 6. Gasteromycetes (
w )

The Gasteromycetes are distinguished from the

Hymeuomycetes by their closed
which open only after the spores are ripe, by the rupture of the
fructifications,
outer hyphal cortex or PERIDIUM. The spores are formed within the fructifications
in an inner mass of tissue termed the GLEBA it contains numerous chambers,
;

which are either filled with loosely interwoven hyphae with lateral branches
terminating in basidia, or their walls are lined with a basidial hymeuium.
The Gasteromycetes are saprophytes, and develop their mycelium in the humus
soil of woods and meadows. Their fructifications, like those of the Hymenomycetes,
arc raised above the surface of the substratum, except in the group of the Hymeno-

gastreae, which possesses subterranean, tuberous fructifications resembling those of

the Tuberaceae.
The fructifications of Sclcrodcrma vulgarc (Fig. 363, 1) have a comparatively
simple structure. They are nearly spherical, usually about 5 cm. thick, and have
a thick, light brown, leathery peridium which finally becomes cracked and
414 BOTANY

ruptured at tlie apex. The gleba is black when ripe, an<l contains numerous
chambers filled with interwoven hyphse which produce lateral, pear-shaped hasidia
with four sessile spores (Fig. 363, 2). This species, which is considered pois<>nou>,
is sometimes mistaken for one of the Truffle Fungi.

The genera Hovista and Lycoperdon (Fig. 363 3) (Pulf-

balls) have also spherical fructifications, which are at first
white, and later of a brown colour. In the last-named
genus they are also stalked, and in the ease of Lycopcnlmi
Bovista may even become half a metre in diameter. The
peridium is forme* 1 of two layers; the outer se para te-
at maturity, while the inner dehisces at the summit.
The hymenial layer of basidia, in the fungi of this group,
chambers of the gleba. The chambers are also
lines the

provided with a fibrous capillitium consisting of brown,

thick-walled, branched hyphie whiclijspring from the walls,
and aid in distributing the spores. The fructifications are
edible while still young and white. When mature they
contain urea which is not elsewhere found in plants.

FIG. 363. 1, Sclerotlernia viilgare, fructification ; -, liasidia of Fit;. 3t'>4.

;
I'hallnit iiy<i/, >-><>.

same. (After Tri.ASNE.) 3, Lycoperdon gemmutinii ; /,,

CH^II /
(After KKOMBHOLZ, A nat.
ii-ininlnxui'. (/, ..'. /, nat. si/i-; .'enlarged.) -i/c.)

In the related genus Geaster (Earth-star) (Fig. 363, 4) the peridia of the nearly
spherical fructifications are also composed of two envelopes. When the dry fruit
dehisces, the outer envelope splits into several stellate segments and the inner
layer of the peridium becomes perforated by an apical opening.
 CRYPTOGAMS 415

The highest development of the fructifications is exhibited by the Phalloideae

82
( of which Phallus impudicus (Stink-horn) is a well-known example.
), This
fungus is usually regarded as poisonous. It was formerly employed in a salve as
a for gout.
remedy Its fructification recalls that of the discomycetous Morchella,
but has quite a different manner of development. A fructification of this
it

species of Phallus is about 15 cm. high. It has a thick, hollow stalk of a white
colour and is perforated with pores or chambers. Surmounting the stalk is a bell-
shaped pileus covered with a brownish-green gleba which, when ripe, is converted
into a slimy mass (Fig. 364). When young the fructification forms a white, egg-
shaped body, and is wholly enveloped by a- double-walled peridium with an inter-
mediate gelatinous layer. Within the PERIDICM (also termed volva) the hyphal
tissue becomes differentiated into the axial stalk and the bell-shaped pileus, carry-

ing the gleba in the form of a mass of hyphal tissue, which contains the chambers
and basidial hymenium. At maturity the stalk becomes enormously elongated,
and pushing through the ruptured peridium raises the pileus with the adhering
gleba high above it. The gleba then deliquesces into a dropping, slimy mass,
which emits a carrion-like stench serving to attract carrion-flies, by whose agency
the spores embedded in it are disseminated.

CLASS XV
Lichenes (Lichens) (. 8M1 )

The Lichens are symbiotic organisms ; they consist of higher

Fungi, chiefly the Ascomycetes, more rarely Basidiomycetes, and uni-
cellular or filamentous Algae (Cyanophyceae or Chlorophyceae), living
in intimate connection, and together forming a compound thallus or
CONSORTIUM. Strictly speaking, both Fungi and Algae should be
classified in their respective orders ; but the Lichens exhibit among
themselves such an agreement in their structure and mode of life,
and have been so evolved as consortia that it is more convenient to
treat them as a separate class .
In the formation of the thallus the algal cells become enveloped
by the mycelium of the fungus in a felted tissue of hyphse (Figs. 365,
366). The fungus derives its nourishment saprophytically from the
organic matter produced by the assimilating alga ; it can also send
haustoria into the algal cells, and so exhaust their contents ( 84 ).
The alga, on the contrary, derives a definite advantage from its
consortism with the fungus, receiving from it inorganic substances
and water, and possibly organic substances also (cf. p. 234). From
the symbiosis entered into by a Lichen Fungus with an Alga, a
dual organism results with a distinctive thallus, of which the form
(influenced by the mode of nutrition of the independently assimilating
Alga) differs greatly from that of other non-symbiotic Eumycetes,
and rather resembles that of the Algae and Liverworts.
The thalli of the Lichens exhibit a great variety of forms.
The simplest Lichens are the FILAMENTOUS, with a filiform
416 BOTAXY

branched thallus consisting of algal filaments interwoven with fungal

hyphae. An example of such a filamentous form is presented by
Ephebe pubescens, which is found growing on damp rocks, forming a
blackish layer.
Another group Is formed by the GELATINOUS Lichens, whose
thallus, xisually foliaceous, is of a gelatinous nature. The algae
inhabiting the thalli of the gelatinous Lichens belong to the families
of the Chroococcaceae and Xostocaceae, whose cell walls are swollen,

forming a gelatinous mass traversed by the hyphae of the fungus.

The genus Collema is a European example of this group.

FIG. 365. Xantlioriu ^mriitinu. 1, Germinating ascdsjHUc (.<//) with branching germ-tube ]i|>lii-<l

to the Chlorococcum cells (a) S, thallus in process of formation sp, two ascospores ; p, Chloro-
; :

coccum cells. By the fusion of the hyphaa in the middle of the mycelium, a pwado-ptNBChj -

matous, cortical layT has br^vm to form. (After BONNIER, from v. TAVEL, x 500.)

In both the filamentous and gelatinous Lichens the algae and

the fungal hyphae are uniformly distributed through the thallus,
which is then said to be unstratified or HOMoiOMEROUS.
The other Lichens have stratified or HETEROMEROUS thalli. The
enclosed algae are usually termed GONIDIA. They are arranged in a
definite GONIDIAL LAYER, covered, externally, by a CORTICAL LAYER,
devoid of algal cells and consisting of a pseudo-parenchyma of closely
woven hyphae (Fig. 366). It is customary to distinguish the three
following forms of heteromerous Lichens. CRUSTACEOUS LICHENS,
in which the thallus has the form of an incrustation adhering closely
to a substratum of rocks or to the soil, which the hyphae to a certain
extent penetrate. FOLIACEOUS LICHENS (Fig. 367), whose flattened,
leaf-like, lobed or deeply-cleft thallus is attached more loosely to the
 CRYPTOGAMS 417

substratum by means of rhizoid-like hyphae (rhizines), springing either

from the middle only or irregularly from the whole under surface.
FRUTICOSE LICHENS (Fig.368) have a filamentous or ribbon-like thallus
branched in a shrub-like manner and attached at the base. They are
either erect or pendulous, or
may sometimes lie free on the
surface of the substratum.
In nature the germinating
spores of 'the Lichen Fungi
appear to be capable of con-
tinuing their further develop-
ment only when they are en-
abled to enter into symbiosis
with the proper gonidia. For n n tT*V V? ^O C^***"- *^~* ^ ** t^*~^-

a few genera of Lichens, how-

ever, it has been determined
that the fungi sometimes exist
FIG. 366. Cetraria islandica. Transverse section
in nature without the pres-
through the thallns ; or, cortical layer of upper
ence of the algae ; it has been surface; ur, of the lower surface; m, medullary

shown that the tropical Lichen, layer containing the green cells of the Alga, Chloro-
corcumhumieolc. (x 272.)
Cwa pavonia (Fig. 374), whose
fungus belongs to the order Hymenomycetes, may produce fructi-
ficationseven when deprived of its alga ; these have a form re-
sembling those of the fungal genus Thelephora. Small thalli have
also been successfully grown from the spores of certain Lichen-

forming Ascomycetes, cultivated without algae and supplied with a

proper nutrient solution.
Many Lichens are able to multiply in a
purely vegetative manner, by means of loosened
pieces of the thallus, which continue their
growth and attach themselves to the sub-
stratum with new rhizines. The majority of
the heteromerous Lichens possess in the forma-
tion of SOUEDIA another means of vegetative
multiplication. In this process, small groups
Fio. 307.- -Xontharia /
on u piece of bark.
of dividing gonidia become closely entwined
(Xat.
si/f. )
with mycelial hyphse, and form small isolated
bodies which, on the rupture of the thallus,
are scattered in great numbers by the wind and give rise to new
Lichens.
The fructifications of the Lichens are produced by the fungi, not
by the algae, which are purely vegetative.

Most Lichens secrete special substances, especially in their cortical layers, among
which numerous acids must lie mentioned ( 85 ) according to STAHL these are useful
;

as a protection against attack 88

by snails ( ).

2 E
418 BOTANY

1. Aseoliehenes

Only a few genera of Lichens have flask -ishaped peritheeia, the fungus lie-
longing to the Pyrenomycetes (Endocarpon, Verrucaria). Most genera produce, as
the ascus- fruit of their fungus, cupular or discoid apotliecia, sessile or somewhat

Fio. 3C.S. Usnea tiarbata. op, Apothecium. Fi. 369. I'etraria island ica. up, Ajiotlii'ciuni.
(Nat. si/e.) (Nat. sixc.)

sunk ill the thallus. In structure they resemble those of the Pezizeae. and
bear on their upper side an hymenium of asci and paraphyses. One of the
commonest species of fruticose Lichens belonging to this group is Usnea burlnitu,
the Beard Lichen, frequently occurring on trees and having large, fringed apotheeia
(Fig. 368). Kocf.Ua tinctoria, found widely distributed on the rocks of the
African coast and East Indies, has an erect, vermi-
form, forked thallus from which litmus and orchil
(orseille) are obtained. Cetraria -islaml'int, Iceland
Moss (Fig. 369), occupies an intermediate position
between the fruticose and foliaceous Lichens. It
has a divided, foliaceous but partially erect thallus,
which is of a light bluish-green or brown colour,
whitish on the under side, and bears the apothecia
obliquely on its margin. This Lichen is found in
Fio. 370. Clndonia. coccifera. mountainous regions in the northern part of the
Scales of primary thallus.
t,
Xortheru Hemisphere, and also at Cape Horn it ;

(Nat. si/..-.) -..11

has an official value as a demulcent. v ,j
Xuiitlwrui
parietina (Fig. 367) may be taken as an example of one of the commonest of
the foliaceous Lichens. The thallus is orange - yellow in colour, and bears
numerous apothecia on its central portions. Graphis script a is a well- known
example of the crustaceous Lichens; its greyish white thallus occurs on the
-

bark of trees, particularly of the Beech, on whose surface the apothecia are
disposed as narrow, black furrows resembling writing. To the crustaceous Lichens
belongs also Sphacrothallia csculenta, growing on rocks in the steppes and deserts
of North Africa and Asia. The thallus falls into small pieces the size of a pea
 CRYPTOGAMS 4] 9

which are scattered by the wind ; they are utilised by the Tartars iu the prepara-
tion of earth-bread.
A peculiar mode of development is exhibited by the genus Cladonia ( 87 ), whose
primary thallus consists of small horizontal scales attached directly to the ground,
from which rises an erect portion, the PODETIUM, of varying form and structure in
the different species. In some cases the podetia are stalked and funnel-shaped,
bearing on the margin or on outgrowths from it knob-like apothecia, which in
C. pxyidata are brown, in C. coccifera (Fig.

370) bright red. In other species the erect

podetia are slender and cylindrical, simple
or forked ;
in C. rangiferina, Reindeer Moss,
which has a world- wide distribution, particu-
larly in the tundras of the North, the podetia vJlr /^*Jk\ .'/

are finely branched (Fig. 371), and bear the

small brown apothecia at the ends of the
branches. The primary thallus of this species
soon disappears.
The ascus -
fructifications (apothecia or

A
A Fio. 3"!-2.f'olli'ina
gonium (c) with
Ki-isjnini.
its
A, Carpo-
trichogyne (t)
FIG. 371. Cladonia rangiferina. A, Sterile ; B, with (X 405). B, Apex of the trichogyne
ascns-fruit at the, ends of the branches. (Nat with the. spermatiuni (.<) attached
si/e.) (x 1125). (After E. BAVR.)

perithecia) of the Lichens originate, as STAHL, and more recently BAUK (**) have
shown, from carpogonia or female sexual organs, which are frequently present in
large numbers on young lobes of the thallus. The carpogonium (Fig. 372) is here
a inulticellular filament, the lower part of which is spirally coiled, while it continues
above into a trichogyne composed of elongated cells, and projecting from the surface
of the thallus. All the cells are uninucleate and communicate with one another by
means of pits. Those of the lower part of the filament contain abundant protoplasm.
Apart from their multicellular nature these structures recall the carpogonia found in
the Florideae. The spermatia, which originate in spermogonia (Fig. 373), are pre-
sumably the male sexual cells. The spermatia develop in different ways ( w ). In
some cases the inner wall of the spermogonium is lined with simple or branched
hyphal branches from the ends of which the spermatia are abstricted (Peltigcra,
420 BOTANY

Parmelia). In other cases the spermogonium is at first filled with a hyphal tissue in
which cavities are formed later and the spermatia arise on very small and thin stalks
from the cells lining the cav-
ities. The em- spermatia.
bedded in a slimy ma. ai
shed from tire spermogoniuni
and conjugate with the ad-
hesive tip of the tricliogyne
(Fig. 372 B). After conjuga-
tion the spermatia appear
empty and their nucleus has

disappeared. When this has

taken place the cells of the
trichogyne collapse, while tin-
cells of the coiled
carpogonium
swell,undergo divisions. an<i
form the ascogonium. From
t He isco< t*n< *u s r
til GIfi r
1 1 *
'

hypha-
J L which bear the asci are
FIG. 373. A*apt)/cMa cilmns. Rii>e Bpennogoniwn. Ihe
dark round bodies within the tlmllus an- the xreen Al.ual produced. Ihe vegetative
cells, (x 192. After GLffcK.) hyphse composing the fructi-
fication and the paraphyses

originate from hyphte which arise below the carpogonium. The fructification may
arise from one or from several carpogonia. The behaviour of the sexual nuclei
requires further investigation. Such carpogonia have been shown to give rise to
the fructifications in a large number of genera. In other genera (Pelliycra,
they are reduced and the trichogyne is
Xo/f rina), wanting, and the reproduction
isapogamous. Spermogonia are, as a rule, not found in these cases, or are, as in
the case of Nephromium, degenerating structures. It has been shown by A.
MOLLER (*), that the spermatia of
Lichens can germinate and produce a
mycelium but this is not inconsistent
;

with their primitively sexual nature.

2. Basidioliehenes (Hymeno-
01
liehenes) ( )

The Hymenolichenes
are represented

by the tropical Cora pavonia, of which

the genera Dictyonema and L<ni.tl<itc<t are
only specially ditt'erentiated forms. The
fungus of this Lichen belongs to the
family Thelephoreae (p. 410) its flat, ;

lobed,and often imbricated fructifications

are also found entirely devoid of Algae.
FIG. 374.-''c,rv< pavoniu. .1, Viewed f'n>m almve
In symbiosis with the unicellular Alga /;. mi below; hym, hytneniiiin.
l'i<
(Xat. si/e.)
Chroococcus, it forms the fructifications of
Cora pavonia (Fig. 374), resembling those of the Thelcphoras with a channelled.
basidialhymenium on the under side. Associated symbiotically, on the other
hand, with filaments of the blue-green Alga ^'c/' / """"", if the Fungus preponderates,
 CRYPTOGAMS 42J

it produces the bracket-like Lichens of the Dictyonema form, found projecting

from the limli.s of trees with a semicircular or
nearly circular thallus, having the
hynienium on the under side. When the shape of the thallus is determined by the
Alga, a Lichen of the Laudatea form occurs as felted patches of tine filaments on
the bark of trees, with the hynienium on the parts of the thallus which are turned
away from the~light.
OFFICIAL. The only representative of the Licliens is Cetraria islandica
KX ISLANDICUS). Sticta pulmouacea is also used in domestic medicine.

9 * 93
II. BRYOPHYTA (MOSSES AND LIVERWORTS) (
- 94
)

The Bryophyta or Muscineae comprise two classes, the Hepaticae

or Liverworts, and the Musd or Mosses. They are distinguished
from the Thallophyta by the characteristic structure of their sexual
organs, ANTHERIDIA and ARCHEGONIA, which are similar to those of
the Pteridophyta, the most highly developed of the Cryptogams.
The Bryophyta and
Pteridophyta are accord-
ingly regarded as having
been derived from a
common ancestor, and,
in contrast to the Thallo-

phyta, they are referred

to collectively as Arche-
goniatae.
The ANTHERIDIA or
male sexual organs are
stalked, ellipsoidal,
spherical, or club-shaped,
with thin walls formed x
MSp|/'
of one layer of cells and
FlG 375.-Jfarefca polymorpha. A Nearly
P n r n i n v Tin m p r n us ripe antiwridium
' ,
1 si
in optical section B, Spermatozoids tix.-d
; p, paraphyses.
Small, Cubical Cells, each with 1 per cent perosmic acid. (A x 90, B x 600.)
of which becomes
divided diagonally into two spermatozoid mother-cells ( ). At maturity
:>

the spermatozoid mother -cells separate and are ejected from the
antheridium, which ruptures at the apex. By the dissolution of the
free as
enveloping walls of the mother-cells the spermatozoids are set
short, slightly twisted filaments, terminating anteriorly in two long cilia.
The ARCHEGONIA are flask-shaped bodies with walls formed of
but one layer of cells ; they are sessile or shortly stalked, sometimes
also somewhat sunk in the tissue, and consist of a dilated ventral

portion and a neck. The ventral portion encloses a large central

cell, the contents of which shortly before maturity divide into the
egg-cell (Fig. 376 A, o) and the ventral -canal-cell (&")'.
The latter
is situated at the base of the neck, just below a central row of neck-

canal-cells (&'). On the maturity of the archegonium, the ventral-

422 BOTANY

and neck-canal-cells become mucilaginous and disorganised. If water

is present, the cells at the apex of the neck separate (B) and the

mucilaginous matter is discharged, and exerts through the diffusion

of certain of its constituents in the water (proteid substances in
96
Marchantia, cane-sugar in the case of Mosses ( )) an attractive
stimulus on the spermatozoids. The spermatozoids, thus directed
toward the neck of the archegonium, traverse it as far as the egg,
into which one sper-
matozoid penetrates.
The water necessary for
the process of fertilisa-
tion is supplied by rain
or dew. After fertilisa-
tion has been accom-
plished, the -
egg cell
divides and gives rise
directly to an embryo
((7),
without first, as is

usually the case in

oospores, undergoing a
period of rest.
The Mosses as well
as the Pteridophytes
multiply also asexually
by means of SPORES
provided with cell-walls
and adapted for dis -
semination through the
air. These two modes
of reproduction, sexual
and asexual, occur in
io.&76.-Mar<'Jtan1iai*>l!iniori.li<t. A, Young, JB, mature arch,-, regular alternation, and
gonium ; C, fertilised archegonium, with dividing egg-cell ; are Confined tO sharply
f, neck-canal-cell ; A"', ventral-canal-cell ; o, egg-cell ; pr, di st i nct generations a :

pseudo-perianth, (x 540.) .

sexual (gametophyte),
provided with sexual organs, and an asexual (sporophyte), which
produces spores. The sexual generation arises from the spore ;

the asexual from the fertilised egg. The number of chromo-

somes in the nuclei of the sporophyte is twice as. great as in the
nuclei of the gametophyte. The double number is acquired in the
fusion of the sexual nuclei, while the reduction to one half takes
place in the division of the spore-mother-cells. THIS ALTERNATION
OF GENERATIONS is characteristic of all Archegoniatae.
In the development of the SEXUAL GENERATION, the unicellular
spore on germinating ruptures its outer coat or EXINE, and gives rise
to a germ-tube. In the case of the Hepaticae, the formation of the
 CRYPTOGAMS 423

plant at once commences, but in most of the Musci a branched,

filamentous PROTONEMA is first produced, composed of cells containing

Jl
FIG. 377. Funarin liyyromctrim. A, Germinating spore ; ex, exine ; /}, protonema ; kn, buds ;

r, rhizoids ; s, spore. (After MULLER-THURGAU ; magnified.)

chlorophyll (Fig. 377 r). The green, filamentous protonema gives

rise to branched, colourless rhizoids (r), which penetrate the sub-
stratum. The MOSS-PLANTS arise from buds developed on the
protonema at the base of the branches.
Protonema and Moss-plant, in spite of an P
the difference in appearance between
them, together represent the sexual
generation. Many Liverworts possess
a thallus consisting of dichotomously
branching lobes, which is attached to
the substratum at its base or on the
under side by means of rhizoids, thus
repeating the vegetative structure of
many Algae (Fig. 379). In other
Hepaticae, on the other hand, and in all
the Musci, there exists a distinct differ-
entiation into Stem and leaves (FigS. pio. 378. Phaseumcuspulntum.ini, Anther.
In no instance, however, idia apex of the
or, archegonia at the
385, 392). ;

r j ,. r bifurcated moss stem '/, leaves ;p, para-

;

are true roots formed consisting of

physes (Aft ,, H( KV1 ISTER x 45 .)
, ,
,

a tissue of cells, but in their stead

rhizoids, consisting of colourless branching filaments, the main function
of which is to attach the plant to the substratum. The Bryophytes,
in this respect, differ essentially from the Pteridophytes, which are
provided with true roots. The stems and leaves of Mosses are also
424 BOTANY PAIIT n

anatomically of a simple structure ; if conducting strands are present,

they are composed merely of simple elongated cells. The sexual
organs are produced on the adult, sexual generation in the thalloid
;

forms, on the dorsal side of the thallus in the cormophytic forms,

;

at the apex of the stem or its branches (Fig. 378).

By the division of the fertilised egg (Fig. 376 6'), a multicellular
embryo is formed, which, by its further development, gives rise to
the second or ASEXUAL GENERATION, represented by the SPOROGONIUM
or the stalked MOSS CAPSULE. The sporogonium, in most cases,
consists of a round or oval capsular receptacle, in whose internal
tissue numerous unicellular spores are produced. At maturity the
capsule opens and sets free the spores. In both the Bryophytes and
Pteridophytes the spores are formed in TETKADS by the twice-repeated
division of the spore-mother-cells, which previously have separated
from one another and become rounded off, and represent the actual
point of commencement of the sexual generation. The spore capsule
has usually a shorter or longer stalk, of which the basal portion, or
foot, remains in the distended venter of the archegonium, and, in
consequence of the overgrowth of the underlying tissue, has the
appearance of being sunk in it. Although the sporogonium constitutes
a distinct asexual generation, it continues throughout its existence
united with the sexual generation, and, like a semi-parasitic plant,
draws from it the nourishment necessary for its development.
As shown by the occasional occurrence of intermediate structures, the antheridia
and archegonia are homologous organs the ventral-canal-cell and the iiffk-ranal-
;

cells of the latter represent gametes which have become funetionless 94 ).

f
Morpho-
logically the sexual organs of Archegoniatae are most closely comparable with the
plurilocular gametangia of the Phaeophyceae, while no close relation can lie tr;n-<-d
with the higher Green Algae. Further, in certain Brown Algae (Dictyotn] a regular
alternation of generations is already present in this case, however, thegametophyte
;

and sporophyte exhibit a similar organisation. The form of the thalloid gami-t<>-
phyte of the lower Liverworts shows the most marked correspondence with tin-
Brown Algae. The sporophyte of Bryophyta has proceeded on other lines <>f
development, but recalls the tetraspore formation in Dictyotn by the tetrad division
of the mother-cells of the asexual spores.

The two very distinct classes of Bryophytes may be briefly charac-

terised as follows :

1. Hepaticae
(Liverworts). The sexual generation, with poorly
developed and generally not distinctly differentiated protonema, is
either a dichotomously divided thallus or is developed as a leafy, and,
with few exceptions dorsiventral shoot. In the majority of Hepaticae,
in addition to spores, the capsule
produces also elaters, sterile cells
which, in their typical development, become greatly elongated and
provided with spiral thickenings (Fig. 382 F). They conduct
nourishment to the developing sporogenous cells, and at maturity,
after the opening of the capsule, serve to separate and scatter the
SECT, i CRYPTOGAMS 425

spores. Only in one order, Anthocerotaceae, does the capsule have a

columella, or an axial mass of sterile cells, which also conduct the
metabolic products to the developing spores.
2. Musci (Mosses). The protonema of the sexual generation is
usually well developed and distinctly defined, and the moss plant is
always segmented into stem and leaves. The leaves are arranged
spirally in polysymmetrical, less frequently in bisymmetrical, rows.
The capsule is always without elaters, but, except in one genus, it

always possesses a columella.

CLASS I

97
Hepaticae (Liverworts) ( )

The Hepaticae are divided, according to the structure of the

sporogonium and the segmentation exhibited by the sexual generation,
into five orders the fiicciaceae, Marchantiaceae, and Anthocerotaceae,
:

comprising exclusively thalloid forms, the Jungermanniaceae, including

both thalloid and dorsiventral foliose forms, and the Calobryaceae,
which are radially constructed foliose forms.

Order 98
1. Ricciaceae ( )

Of all the Hepaticae, this order exhibits the simplest structure. The genus Iticcia

belongs to this order ;

its
clichotomously-lobed or cleft thallus forms small rosettes,
and grows on damp or marshy soil, llicria tmtans is found floating, like Duckweed,
on the surface of stagnant water.
Riccia fiuitans, on the other ham I.
lives wholly submerged, and has

narrow, more profusely branching,

thalloid segments (Fig. 379); it can,
however, grow on marshy soil, and
then forms flat rosettes. The Riecias
are provided with fine rhizoids
springing from the Under side of
the thallus, and possess, in addi-
tion, arow of transversely disposed ,, I(] y!>\-Ricda fluit ,>.x. floating form
J,.Sul.i.i.>rt,"-<l ;

ventral scales, consisting of a single /;, land form. (Nat. siw. K, after GOEBEL.)

layer of cells, which also function

in the absorption of nourishment. Both organs are wanting in the submerged
form of Riccia fluitans.
The antheridia and archegonia are sunk in the surface of the upper side of the
thallus. From the fertilised egg-cell is developed a spherical sporogonium which
has no stalk. The wall of the sporogonium consists of a single layer of cells it ;

becomes disorganised during the ripening of the spores, which are eventually set
free by the rupture and disintegration of tin; venter and the surrounding cells of the
thallus.
426 BOTANY

Order 2. Marehantiaeeae ("")

Tlic Liverworts included in this order are much more highly organised, and in
many genera they have a decided complicated structure. .V" /<// />////" //.//v//"-//'/"'.
found growing on damp soil, may serve as an example. It forms a flat deeply-

lohed, dichotomously-branched thalltis, about two centimetres wide, and having an

inconspicuous midrib (Fig. 381 A, Fig. 382 A). From the under side of the thallus
spring unicellular rhizoids, of which some have smooth walls and serve mainly
to attach the thallus, while others have conical thickenings projecting into the cell-
cavity these peg-rhizoids serve for water conduction.
;
The thallus is provided
also with ventral scales, consisting of a single layer of cells. The dorsiventrality
of the thallus is further shown by its complicated anatomical structure. With the

Fio. 380. Mitii-li'iiitni i*ilyiii<iriili<i. A-C,

Successive stages in the formation of a
gemma; st, stalk-cell; /). surface view;
K, transverse section of a gemma B, ; Fl<;. 3)S1. Man-kit iiliii jxi///w(i/^(7i". .1, A mule
point of attachment to stalk oil cells ; ,
:
plant, with antheridiophores and gemma-cups >>
r,colourless cells with granular contents. (nut. si/e) /;, section of young antlieriilio])hoi-e
; :

from which the rhizoids will develop. a. antheridia : /, thallus; s, ventral seal-
(After KNY, A-C x 275 D-K x 65.) ; rhizoids. (Somewhat magnitled.)

naked eye it may be seen that the Upper surface of the thallus isdivided into small
rhombic areas. Each area is perforated by a central air-pore leading into a corre-
sponding air-chamber immediately below (Fig. 160 A, B). The lateral walls of
the air-chambers determine the configuration of the rhombic areas. The air-
pore in the roofing wall of each chamber is in the form of a short canal, bounded
by a wall formed of several tiers of cells, each tier comprising four cells. Numerous
short filaments, consisting of rows of nearly spherical cells containing chlorophyll
grains, project from the floor of the air-chambers and perform the functi<m> "I
assimilating tissue. Chlorophyll grains are found also in the walls and roof of the
chambers, but only in small numbers. The air-chambers represent depressions in
the outer surface which have become roofed over by the more rapid growth of the
adjacent epidermal cells. The intensity of the illumination exercises a great
influence on the formation of the air chambers ; when the illumination is very weak
they may not occur at all. The epidermis on the under side of the thallus is

formed of one layer of cells. The tissue below the air-chamber layer is devoid of
chlorophyll, and consists of large parenchymatous cells, which serve as storage cells.
 CRYPTOGAMS 427

Small cup-shaped outgrowths, with toothed margins, the gemmiferous receptacles

or gemma-cups, are generally found situated on the upper surface of the thallus over

Fni. :iv_>. Mni-i'liniitiii jK>l)/iniii-iihii. A, A female, plant, with four archegoniophores ot'diflerent a^es ;

ft, gemma-cups (nat. size). B, under side of receptacle;** rays: 'i. sheath ;.-;/ sj>orogonium(x3).
C, half of a receptacle, divided Longitudinally (x5). D, longitudinal .section of a young si>oro-
Konitim ; npf, the foot; x/'- spni-d^ennus tissue /. wall, and h neck, of
: /.'. wall of capsule :

arche^oniiini ;<, pBeado-periantb ( X 70). K, ruptured sporoj,'<miuin

; capsule; SJKUVS and ; /,- .-.

elaters ji, |isendo-]>eriarith ; c, archegonial wall (x 10).

; F nn'elater. G, ripe sj)ores (x 315).
H, nerminatinn s]Kii-e (x) c/.. protoneina (/,).!j;enii-<lisc. with the apical cell r and rhizoid rh
; ;

(X 100). (<: !: after His< HoFK ; /;. Ii. J-'-H after KSY.)

the midribs (Fig. 381 b). These contain a number of stalked gemmae, flat, biscuit-
shaped bodies of a green colour. The gemmae arise by the protrusion and repeated
division of a single epidermal cell (Fig. 380); at maturity they become detached
from the stalk (at x, Fig. 380 D). They are provided with two growing points.
428 BOTANY I-AUT n

one at each of the marginal constrictions from which their further development
into new jilants proceeds. On cross-section (E) they are seen to lie composed of
several layers of cells some of the cells are lilled with oil globules (7), o), while
;

from other colourless cells rhizoids develop. Cells containing oil are also present
in the mature thallus, and are of frequent occurrence in all the Hepaticae. By
means of the abundantly developed genuine Murr/mntin is enabled to multiply
vegetatively to an enormous extent.
The sexual organs, antheridia and archegonia, are borne on special erect branches
of the thallus. The reproductive branches, which are contracted below into a
stalk, expand above into a profusely branched upper portion. In this species,
which is dioecious, the antheridia and archegonia develop on dilferent plants.
The branches producing the male organs terminate in lobed discs, which bear the
antheridia on their upper sides in flask-shaped depressions, each containing an
antheridium (Fig. 381 B). The depressions, into each of which a narrow canal
leads, are separated from each other by tissue provided with air-chambers. (The
structure of the antheridia and sperniatozoids is illustrated by Fig. 375 ami the
accompanying description. )
The female branches terminate each in a nine-rayed disc (Fig. 382 A). The
upper surface of the disc, between the rays, becomes displaced in the processor
growth, and, as the archegonia are borne on these portions, they seem to arise from
the under side of the disc. The archegonia are disposed in radial rows between the
rays, each row being developed in a toothed lamella or sheath (Fig. 382 />', C', li ;

for structure of the archegonia see Fig 376 and description .

The fertilised egg-cell gives rise to a multicellular embryo (Fig. 376 C), and
this, by further division and progressive differentiation, develops into a stalked
oval SPOROOONIUM. The capsule of the sporogonium is provided with a wall con-
sisting of one layer of cells, the walls of which have thickened bands, and ruptures
at the apex to let free the spherical spores. The ELATEKS, or elongated, spirally
thickened cells formed in the capsules, between the spores, by the prolongation of
definite cells, are characteristic of the Marchantias and most of the Liverworts,
except the Ricciaceae. The elaters are discharged from the ruptured capsule, together
with the spores, and serve for their dispersion in the same way as the capillitium
of the Myxomycetes (Fig. 382, E, F, G). The ripe capsule, before the elongation
of the stalk, remains enclosed in the archegonium wall D, mv), which, for a time,
keeps pace in its growth with that of the capsule. As the stalk elongate-, the
archegonium wall or calyptra is broken through and remains behind, as a sheath,
at the base of the sporogonium (E, c). The capsule is surrounded also by the
pseudo-perianth, an open sac-like envelope which grows, before fertilisation, out of
the short stalk of the archegonium (Fig. 376 C, pr Fig. 382 D, E, p).
;

Marchantia was formerly used in the treatment of diseases of the liver this ;

fact explains the origin of the name Liverwort.

Order 3. Anthoeerotaeeae
The few forms included in this order have an irregular, disc-shaped thallus,
which is firmly anchored to the soil by means of rhizoids. The cells of the thallus
contain, in contrast to those of other Hryophyta, a single large chloroplast. The
antheridia arise, in groups of two to four, by the division of a cell lyhig below the
epidermis ; they remain enclosed in cavities beneath the upper surface of the
thallus until maturity. The archegonia are sunk in the upper surface of the
thallus ;
after fertilisation they become covered over by a many-layered wall
 CRYPTOGAMS 429

formed by the overarching growth of the adjoining tissue. This enveloping wall
isafterwards ruptured by the elongating capsule, and forms a sheath at its base.
The sporogonium consists of a swollen foot and a long, pod-shaped capsule it has ;

no stalk. The superficial cells of the foot grow out into

The capsule splits longitudinally
rhizoid-like papilla-.
into two valves, and has a central hair-like columella
formed of a few rows of sterile cells (Fig. 383). The
columella does not extend to the apex of the capsule,
but is surmounted by a narrow layer of sporogenous cells.
Elaters also occur they are multicellular, variously
;

shaped, and often forked. The sporogonia, unlike those

of all other Hepaticae, do not ripen simultaneously
throughout their whole length, but from the tips down-
wards, and continue to elongate by basal growth after
('merging from the archegonia. The wall of the sporo- FIG. 383. Anthoceros Jam's.
sp, SiK)rogonium_; colu-
gonium possesses stomata, which do not occur in other
c,
ini'lla. (Nat. sixi>.)
Liverworts ; present in its cells.
chlorophyll is

On the under side of the thallus, slit-like openings, formed by the separation
of the cells, lead into cavities filled with mucilage. Nostoc filaments penetrate
into these cavities, and develop into endophytic colonies.

Order 4. Jungermanniaeeae
These are usually small forms which grow on the ground or on tree-trunks and
in the tropics on the surface of living leaves. In the simplest forms of this order
the thallus is broadly lobed, similar to that of Marchantia (e.g. Pellia epiphylla,
frequently found on damp ground) ;

or, like that of liiccia Jiuitans, it is

narrow and ribbon-shaped, and at the
same time profusely branched (e.g.
.}/</ -.i/cr-ia furcata, Fig. 163). In the

Ki. 3S4. lutia iHisiliii. .-.-, Sporogoninm ;

Fia. 385. Plagiocliila asphniiiules.
r, rhizoids. (x 2.) gonium. (Nat. size.)

again, the broad, deeply lobed thallus has an evident midrib,

-
other forms,
and its margins, as in the case of Blasia pusilla (Fig. 384), exhibit an in-
cipient segmentation into leaf-like members. The majority of Jungerman-
430 BOTANY

niaceae, however, show a distinct segmentation into stem and leaflets. The
latter consist of one layer of cells without a midrib, and are inserted with ob-
liquely directed laminse in two rows on each flank of the stem. Some species
(e.g. Frullania Tamarisci, a delicately branched Liverwort of
a brownish colour

occurring on rocks and tree-trunks) have also a ventral row of small scale-like
leaves or amphigastria (Fig. 386 a). The dorsal leaves are frequently divided into an
upper and lower lobe. In species growing in dry places, like Frullania ?'///"//*</,
the lower lobe is modified into a sac, and serves as a capillary water-reservoir. The
leaves regularly overlap each other they are then said to be succubous, when the
;

posterior edges of the leaves are overlapped by the anterior edges of those next
below (Frullania, Fig. 386), or in-
cubous, if the posterior edges of the
leaves overlap the anterior edge* of
the leaves next below (Plagwchilii,
Fig. 385).
The branching stems of the foliose

'ws

FIG. 380.Part of a shoot of Frullaiiia

Tamarisci, seen from below. >, Dorsal

leaves with the lower lobes (u) modified FIG. 387. Haplormitrium Hookeri. a, origin of
as water-sacs ; <i, amphiKastriiuii. ( x a new shoot r, rhizome ; o, lower limit-of

36.) tin- iii-rial shoot. (After OOTTISCHE.)

Jungermamiiaceae are either prostrate or partially erect, and in consequence of the

manner in which the leaves develop, present a distinctly dorsiventral appearance.
The long-stalked sporogonium is also characteristic of this order ;
it is already
fully developed before it is pushed through the apex of the archegonial wall by
the elongating stalk. It has a spherical capsule which on rupturing splits into
four valves (Figs. 384, 385). No columella is formed in the capsule but in ;

addition to spores it always produces elaters, which, by their movements while

drying, scatter the spores. The stalk of the capsule is delicate. The wall of the
capsule (usually two or several cells thick) consists of cells with annular or band-
like thickenings, or the walls are uniformly thickened with the exception of the
outermost walls. Dehiscence is dependent on the cohesive power of the water in
these cells causing the outer walls to bulge into the cavity.
According to the position of the sexual organs and sporogonium the Junger-
manniaceae are divided into two groups. In the Anakrogynae the apex is nut
used up in the formation of the archegonia, and the sporogonia are situated on
SHOT, i CRYPTOGAMS 431

the dorsal surface and are surrounded by a sheath-like outgrowth of the thallus
forming an involucre. To this group belong the thalloid forms (Pellia, Metz-
ycria) and others showing a transition to the frondose forms (Blasia). In the
Akrogyiuie, on the other hand, the archegonia and the sporogoniurn stand at
the end of the main stem or of a branch and are surrounded by a perianth
formed of modified leaves. To this group belong the dorsiventral leafy forms,
c.;/. I'linjiiiclrila (Fig. 385), Frullania, and Jungcrmannia, a genus with numerous
species.

10
Order 5. Calobryaeeae ( )

This order contains only two genera, Calobryum occurring in the tropics and
Jfaplomitrium. The single species of the latter genus H. Hookeri (Fig. 387)
occurs in Europe, and possibly is a survival of preglacial Liverworts. The
Calobryaeeae differ from all other Liverworts in the radial construction of their
shoot, which bears three rows of leaves. The sexual organs form terminal groups
in Culibryum, in Haplomitrium they occur between the upper leaves.

CLASS II

101
Musci (Mosses) ( )

The profusely branched protonema of the Mosses appears to the

naked eye as a felted growth of fine, green filaments (Fig. 377).
The young moss plants are developed on the protonema as small
buds which arise as protrusions of cells of the filament, usually from
the basal cell of one of the branches. The protrusion is cut off by a
transverse septum, and after the separation of one or two stalk-cells
the three-sided pyramidal apical cell of the moss plant is delimited
102
in the enlarged terminal cell ( ).
The moss plants are always
differentiated into stem and leaf. The Mosses may be readily dis-
tinguished from the foliose Jungermanniaceae by the spiral arrange-
ment of their small leaves, which are rarely arranged in two rows.
In Mosses which have prostrate stems, the leaves, although arranged
spirally, frequently assume
a somewhat outspread position, and all
face one way, so that in such cases a distinction between an upper
and a lower side is manifested, but in a manner different from that
of the Liverworts.

The STEM OF THE Moss PLANT is formed of cells which become gradually
smaller and thicker-walled towards the periphery. In the stems of many genera
(e.y. Mnium, Fig. 161) there is found a central, axial strand consisting of
elongated, conducting cells with narrow lumina. These strands are not as highly
differentiated as the vascular bundles of Pteridophytes. They do not occur in the
Sphagnaceae or Bog-mosses which grow in swampy places. The stems of the Sphag-
naceae show a peculiar development of the outer cortical layers (Fig. 388 C). The
devoid of protoplasm, and are in communication witli each
cells in these layers are
other and the atmosphere by means of large, open pores ; to secure rigidity, they
432 BOTANY

are also provided with spirally thickened walls. They have a remarkable power
of capillary absorption, and serve as reservoirs for storing and conducting water.
The M-:AVF.S of the true Mosses have, as a rule, a very simple structure. They
consist usually of a single' layer of polygonal cells containing chloroplasts (Fig. 63
and Fig. 100, Funaria), and are generally provided with a conducting bundle of
elongated cells. The leaves of the Bog Mosses (Sphagnaceae) have no bum lies,
and instead are supplied with capillary cells for the absorption and storage of
water. These cells are devoid of protoplasm, and are similar to those in the
periphery of the stem, but larger and more elongated their walls, which are ;

perforated, are strengthened by transverse thickening bands (Fig. 388 A, B).

Between them are other elongated, reticulately united cells containing chloro-

c
Fi<;. 388.^, Surface view of :i portion of a leaf of Sphagnum cymbifolium (x 300). li, part of a
transverse section of a leaf of Sphagnum fimbriatum a, evil containing chlorophyll:
;

w, capillary cell v, thickening hands

; 7, j>ore. (*, part of a transverse section of the stem of
;

Sphagnum cymbifolium ; c, central cells ; sk, sclerenchymatous cortical cells ; w, capillary cells
with i>ores (1) ; e, epidermis, (x 120.)

plasts. A similar differentiation of the leaf cells occurs in a few other Mosses
(e.g.Leucobryum vulgare).
A more complicated structure of the leaves resulting from their adaptation to
the absorption of water is exhibited by Polytrichum commuiw. In this Moss tin-
leaves develop on their upper surface numerous, crowded, vertical lamellae, one
cell thick ; these contain chlorophyll and function as assimilatory tissue, while
the spaces between the lamella' serve as reservoirs for the storage of water. In a
dry atmosphere the leaves fold together, and thus protect the delicate lamella*
from excessive transpiration.
The RHIZOIDS (Fig. 395), each of which consists of a branched filament of cells,
spring from the base of the stem. In structure they resemble the protonema, into
which they sometimes become converted, and then give rise to new Moss plants.

The SEXUAL ORGANS are always borne in groups at the apices

either of the main axes or of small, lateral branches, surrounded by
their upper leaves ;
each group with its involucral leaves constituting
 CRYPTOGAMS 433

a receptacle. The antheridial and archegonial receptacles are some-

times inappropriately referred to as moss flowers, but they have
nothing in common with the true flowers of vascular plants ; the
involucral leaves, which frequently have a distinctive structure, are
also known as the PERICH^ETIUM (Fig. 395). Between the sexual
organs there are usually present a number of multicellular hairs or

paraphyses. The moss plants may be monoecious, in which case

em

FKI. 389. Sphagnum fimbriatuni: A, A shoot with four ripe sporogonia. Sphagnum antti/olium
B, Archegonium with the multicellnlar embryo of the sporogonium em; C, a young sporo-
gonium in longitudinal section ps, pseuuopodium ca, archegonial wall or calyptra ah, neck of
; ; ;

archegonium spf, foot of sporogonium k, capsule

; ; co, columella
; ;spo, spore-sac with spores ;

K, ruptured antheridium with escaping spermatozoids F, single spermatozoid, highly

;

magnified. Sphagnum squarrosum : D, A lateral shoot with a terminal sporogonium ca, ;

ruptured calyptra d, operculum. (After W. P. SCHIMPER A, nat. size the other figures
; ; ;

magnified.)

both kinds of sexual organs are borne on the same plant either in
the same or different receptacles or dioecious, and then the antheridia
;

and archegonia arise on different plants.

The SPOROGONIUM of the Mosses ( 103 ) develops a capsule with
an axial COLUMELLA consisting of sterile tissue. The spore-sac
surrounds the columella, which accumulates food material and water
for the developing spores. Elaters are never formed. In the young
sporogonium outside the spore-sac, a well-developed assimilating
tissue is present; this is bounded by water-storage tissue and an
epidermis. In most Mosses stomata are found on the lower part of
2 F
434 BOTANY

the capsule. Distinctive variations in the mode of development and

structure of the capsules are exhibited by the four orders of the

Fio. 3pO. Andreaea petrophila. ps, Pseudopodium;

.S;>/, foot ; k, capsule ; c, calyptra. (x 12.)

Fio. 391. Ephemerum serratum. p, Protonema ; FIG. 3U2. Pdytrichum commune, rh, Hhi-
ft, foliage-leaf ; sporogonium c, calyptra
s, ; ; zoids ; s, seta ; c, calyptra ; ap, apophysis ;

rh, rhizoids. (After W. P. SCIIIHPER, x 200.) d, operculum. (Nat. size.)

Musci Sphagnaceae, Andreaeaceae, Phascaceae, and Bryinae.

: The
Sphagnaceae and Andreaeaceae are nearest to the Liverworts.

Order 104
1 .
Sphagnaceae ( )

The order of the Sphagnaceae, or Bog Mosses, includes only a single genus,
Sphagnum, containing many species. The Bog Mosses grow in swampy places,
 CRYPTOGAMS 435

and form large tussocks saturated with water. The upper extremities of the
stems continue their growth from year to year, while the lower portions die
away and become eventually converted into peat.
Of the numerous lateral branches arising from each
of the shoots, some grow upwards and form the
apical tufts or heads at the summits of the stems ;

others, which are more elongated and flagcllifonn

in shape, turn downwards and envelop the lower
portions of the stem (Fig. 389 A). Every year one
branch below the apex develops as strongly as the
mother shoot, so that the stem thus becomes falsely
bifurcated. By the gradual death of the stem from
below upwards the daughter shoots become separ-
ated from it, and form independent plants. Special
branches of the tufted heads are distinguishable by
.their different structure and colour on these the
;

sexual organs are produced. The male branches

give rise, near the leaves, to spherical stalked

antheridia, which open at the apices by means of

valves these bend back and let free the spirally FIG. 393. Schistostega osiwindacea.
;
Sterile B, fertile plant, (x 5.)
twisted spermatozoids (Fig. 389 E, F). The arche- A, ;

gonia are borne at the tips of the female branches. The sporogouium develops a
short stalk with an expanded foot (B, 0), but remains for a time enclosed by the

FIG. 395. AInlum undulatum. Orthotropous

shoot terminating in a male receptacle sur-
rounded by involucral leaves. The lateral
FIG. 394. Hypnum purum. (Nat. size.) shoots are plagiotropous. (After GOEEEL.)

archegonial wall or calyptra. Upon the rupture of the archegonium, the calyptra
persists as in the Hepaticae at the base of the sporogonium. The capsule is spherical
and has a dome-shaped columella, which in turn is overarched by a hemispherical
436 BOTANY PART n

spore-sac (spo) ; it possesses an operculum, but no peristome. The ripe sporogoniuin,

like that of Andreaea, is borne upon a prolongation of the stem axis, the pseudo-

podium, which is expanded at the top to receive the foot of the stalk. Of the
peculiar structure of the leaves and stem cortex a description has already been
given above. The protonema of the Sphagnaceae is in some respects peculiar.
Only a short filament is formed on the germination of the spore, the protonema
broadening out almost at once into a flat structure on which the young moss
plants arise.

Order 2. Andreaeaeeae
The Andreaeaeeae comprise only the one genus Andreaea, small, brownish
caespitose Mosses growing on rocks. The sporogonium is also terminal in this
order. The capsule, at first provided
with a calyptra, splits into four longitudinal
valves (schizocarpous), which remain united at the base and apex (Fig. 390). The
stalk is short, and is expanded at the base into a foot (Spf), which in turn is
borne on a pseudopodium (ps), a stalk-like prolongation of the stem resulting from
itselongation after the fertilisation of the archegonium.

Order 3. Phascaceae
To the Phascaceae (Cleistocarpae) belong small Mosses with few leaves growing
on the soil they retain their filamentous protonemata until the capsules are ripe,
;

and have the simplest structure of all the Mosses (Fig. 391). The capsule is
terminal and has only a short stalk it is protected by a calyptra.
;
It does not

open with a lid, but the spores are set free by the decay of its walls.

105
Order 4. Bryinae( )

In this order (termed also Stegocarpae), which includes the majority of all the
true Mosses, the moss fruit attains its most complicated structure. The ripe
SPOROGONIUM, developed from the fertilised egg, consists of a long stalk, the SETA
(Fig. 392 s), with a FOOT at its base, sunk in the tissue of the mother plant, and
of a CAPSULE, which in its young stages is surmounted by a hood or CALYPTRA.
The calyptra is thrown otf before the spores are ripe. It consists of one or two
layers of elongated cells, and originally formed part of the wall of the archegonium
which, at first, enclosed the embryo, growing in size as it grew, until, finally
ruptured by the elongation of the seta, it was carried up as a cap, covering the
capsule. In the Liverworts the calyptra is, on the contrary, always pierced by
the elongating sporogonium, and forms a sheath at its base. The upper part of
the seta, where it joins the capsule, is termed the APOPHYSIS. In Alnium (Fig.
398 A, ap) it is scarcely distinguishable, but in Polytrichum commune it has tin:
form of a swollen ring-like protuberance (Fig. 392, ap), while in species of
Splachnum it dilates into a large collar-like structure of a yellow or red colour.
The upper part of the capsule becomes converted into a lid or operculum which is
sometimes drawn out into a projecting tip. At the margin of the operculum a
narrow zone of epidermal cells termed the ring or ANNTLUS becomes specially
differentiated. The cells of the aunulus contain mucilage, and by their expansion
at maturity assist in throwing off the lid. In most stegocarpous Mosses the
mouth of the dehisced capsule bears a fringe, the PEKISTOME, consisting usually
of tooth-like appendages.
SECT. I CRYPTOGAMS 437

The peristome of Mnium hornum (Fig. 398) is double the outer peristonie ;

is formed of 16 pointed, transversely striped teeth inserted on the inner margin

of the wall of the capsule. The inner peristome lies just within the outer, and
consists of cilia-like appendages, which are ribbed on the inner side and thus
appear transversely striped they coalesce at their base into a continuous membrane.
;

Two cilia of the inner peristome are always situated between each two teeth of
the outer row. The cilia facilitate the dissemination of the spores by their
hygroscopic movements.
The teeth and cilia of the peristome are formed in this instance of thickened

ap

Fio. 397. Mnium hornum. Transverse section

through the wall of the capsule in the region
FIG. 396. Mnium hornum. Median .longi-
of the ring, a, Cells of the ring ; 1-k, succes-
tudinal section of aihalf-ripelsporogonium. sive cell layers with the thickened masses of
o Operculum p, peristome ; c, columella
; ;
the peristome, d', d"; d'", transverse pro-
3,spore-sac containing the spores ; i, air- jecting ribs ; c, the .coalesced cilia, (x
space ; ap, apophysis gt, stomata.
; ( x 18.) 240.)

cells next to the operculum (Fig.

portions of the opposite walls of a single layer of
397), the teeth from portions of the external wall, and the cilia from portions of
the internal walls of the same layer. On the opening of the capsule the un-
thickened portions of this layer break away and the teeth and cilia split apart.
The transversely ribbed markings on their surface indicate the position of the
former transverse walls.
The structure of the varies greatly within the Bryinae.
peristome By its
peculiar form and hygroscopic movements the peristome
causes a gradual dis-
semination of the spores from the capsule.
Variations in the form of the capsule, peristome, operculum, and calyptra afford
the most important means of distinguishing the different genera. The Bryinae
438 BOTANY

are divided into two sub-orders, according to the position of the archegonia or of
the sporogonia developed from them.
(a) Bryinae acrocarpae'. The archegonia, and consequently the sporogonia, are
terminal on the main axis. Mnium undulatum (Fig. 395) and hornum, Poly-
trichum commune (Fig. 392), and Funaria hygrometrica are common examples.
Schistostega osmundacea, a moss living in caves, has fertile shoots, which have
spirally arranged leaves and bear stalked capsules devoid of peristomes, and also
other shoots that are sterile, with two rows of leaves (Fig. 393 A, B). The
protonema of this species is peculiarly constructed and gives out an emerald
phosphorescent light. (Of. Figs. 172, 214.)
(b) Bryinae pleurocarpae. The growth of the main axis is unlimited, and the

Fio. 308. Mnium hornum. A, Capsule with upper portion of seta ap, apophysis p, peristome ;
; ;

d, the separated operculnm. B, Three teeth of the outer peristome seen from the outside ; an,
annulus. C, inner peristome seen from 'the inside ; w, broader cilia ; h, narrower cilia.
(A x 4 ; B, C x 60.)

archegonia with their sporogonia arise on short, lateral branches (Fig. 394). In
this group are included numerous, usually profusely branched species of large
Mosses belonging to the genera Hylocomium, Neckera, and Hypnum, which are
among the most conspicuous mosses of our woods, and also the submerged Water
Moss, Fontinalis antipyretica.

III. PTERIDOPHYTA (VASCULAR CRYPTOGAMS) (

,

The Pteridophytes include the Ferns, Water-Ferns, Horse-tails, and

Club Mosses, and represent the most highly developed Cryptogams.
In the development of the plants forming this group, as in the Bryo-
phyta, a distinct alternation of generations is exhibited. The first
generation, the sexual, bears the antheridia and. archegonia; the
second, the asexual, develops from the fertilised egg and produces
asexual, unicellular spores. On germination the spores in turn give
rise to a sexual generation.
SECT. 1 CRYPTOGAMS 439

The SEXUAL GENERATION is termed the PROTHALLIUM or GAMETO-

PHYTE. It never reaches any great size, being at most a few centi-
metres in diameter; in
some forms it resembles
in appearance a simple,
thalloid Liverwort; it then
consists of a small green
thallus, attached to the
soilby rhizoids springing
from the under side (Fig.
399 A). At other times
the prothallium is
branched and filament-
ous sometimes it is a
;

tuberous, colourless mass

of tissue, partially or
wholly buried in the
ground, and leading a
Fie. 3'.H>. Aaiiiilniin jUiji m.. .1, I'rotlmlliiini seen from
saprophytic existence, lic-ln\v archegonia an, antheridia rh, rhizoids ; B,
; in-, ; ;

while in certain other prothallium with young Fern attached to it by its foot ;
divisions of the Pterido- b, the first leaf; ir, the primary root, (x circa 8.)

phyta it undergoes reduc-

tion and remains more or less completely enclosed within the spore.
On the prothallia arise the sexual organs, antheridia (Figs. 404, 411),
producing numerous
ciliate, usually spiral
spermatozoids, and
archegonia (Figs. 405,
412), in each of which
is a single egg-cell.
As in the Mosses the
presence of water is
necessary for fertilisa-
tion. The spermato-
zoids are induced to
direct their motion
toward the archegonia
FIG. 400. -A, rt,,-;* >,/;"//. emhry,. freed from th. archegonium, by the excretion from
in longitudinal section (after KIENITZ-(JERLOKF) basal wall
:
1, ;
the latter of a sub-
II,transverse wall dividing the egg-cell into quadrants rudi- ;

ment of the foot/, of the stem s. of the lirst leaf 6, of the root stance which diffuses
w ; H, section of a farther-developed embryo of Pteris aquilina into the surrounding
(after HOFMEISTER); /, foot still emlwdded in the enlarged water. In Ferns, Sal-
venter of the archegonium aw ; pr, prothallium. (Magnified.)
vinia, Equisetum, Sela-
107
ginella, and Isoetes, this substance is malic acid or one of its salts ( ).

After fertilisation" the egg-cell develops into a multicellular

embryo, which becomes the asexual generation, as in the Bryophyta.

440 BOTANY

The Bryophyta and Pteridophyta have accordingly been classed

together as Embryophyta by ENGLER, and termed Embryophyta
zoidiogama because the male cells
are developed as spermatozoids.
The asexual generation or sporo-
phyte is represented by a plant
possessing a highly differentiated
internal structure, and externally
segmented into stem, leaves, and
roots. In the majority of Pterido-
phytes (Ferns, Equisetum), the fertil-
ised egg-cell, while still in the arche-

gonium, surrounds itself with a cell

wall and undergoes division, first into
two cells, by the formation of a
Jt
transverse or basal wall, and then
into octants by two walls at right
FIG. 401. Transverse section of the rhizome
of Pteris aquilina. s, Concentric vascular angles to each other and to the basal
peripheral bundles I, sclerenchymatous
; wall. By the further division of
plates ; Ip, zone of sclerenchymatous.flbres; a small mass of
these eight cells
JJ, cortex ; e, epidermis, (x 7.)
tissue is formed, and from this are
developed the stem apex, the first leaf, the primary root, and an

FIG. 402. Transverse section of stem of Lycopodium cornea nntum. rp, Epidermis i-t, vi,pp, outer,
;

inner,and innermost parts of the primary cortex, surrounding the central cylinder formed
by the coalescence of concentric bundles; sc, scalariform trachcides s}>, annular and spiral
;

tracheides v, phloem, (x 26.)

;

organ peculiar to the Pteridophytes, the so-called FOOT (Fig. 400 /).
The foot is a mass of tissue, by means of which the young embryo
 CRYPTOGAMS 441

remains attached to the parent prothallium and absorbs nourishment

from it, until, by the development of its own roots and leaves, it is
able to nourish itself independently. The prothallium then usually
dies. The stem developed from the embryonic rudiment may be
either simple or bifurcated, erect or prostrate ; it branches without
reference to the leaves, which are arranged spirally or in whorls,
or occupy a dorsiventral position. Instead of rhizoids, true roots
are produced, as in the Phanerogams (cf. Fig. 167). The leaves also
correspond in structure with those of the Phanerogams. Stems,
leaves, and root are traversed by well-differentiated vascular bundles,
and the Pteridophytes are, in consequence, designated Vascular
Cryptogams. The bundles of the great majority of Pteridophytes are
constructed on the concentric type (cf. Fig. 401,
and Fig. 125, p. 115).
Secondary growth in thickness, resulting from the activity of a special
cambium, occurs only occasionally in existing forms, but it was char-
acteristic of the stems of certain extinctgroups of Pteridophytes.
The SPORES are produced vegetatively in special receptacles termed
SPORANGIA, which occur on the asexual generation, either on the
leaves, or less frequently on the stems in the axils of the leaves.
The leaves which bear the sporangia are termed SPOROPHYLLS. The
sporangium consists of a wall enclosing the sporogenous tissue, the
cells of which, becoming rounded off and separated from each other
as spore -mother -cells, give rise each to four tetrahedral spores
(spore-tetrads). The cells of the innermost layer of the sporangial
wall are rich in protoplasm, and constitute the TAPETUM. This
layer persists in the Lycopodineae, but in the case of the Ferns and
Equisetineae the walls of the tapetal layer become dissolved. In
the course of the development of a sporangium the tapetal cells
then wander in between the spore-mother-cells, so that the spores
eventually lie embedded in a mucilaginous protoplasmic mass, the
PERIPLASM, from which they derive nourishment. The wall of the
mature sporangium is formed of one or a number of layers of cells.
The unicellular spores have walls composed of several coats.
The spores of the majority of the Pteridophytes are of one
kind, and give rise on germination to a prothallium, which produces
both antheridia and archegonia. In certain cases, however, the
prothallia are dioecious. This separation of the sexes extends in
some groups even to the spores, which, as MACROSPORES, developed
in MACROSPORANGIA, give rise only to female prothallia ; or as MICRO-
SPORES, which are produced in MICROSPORANGIA, develop similarly
only male prothallia. In accordance with this difference in the spores,
a distinction may be made between the HOMOSPOROUS and HETERO-
SPOROUS forms of the same group ; but this distinction has no
systematic value in denning the different groups themselves, as it has
arisen in groups in other respects quite distinct.
Compared with the Bryophyta, the asexual cormophytic generation
442 BOTANY PART n

of the Pteridophytes corresponds to the sporogonium, the prothallium

on the other hand to' the moss plant with its protonema ; although
both groups may have originated phylogenetically from a common
ancestor, they have followed altogether different directions in the
course of their further development. The correspondence in the
structure of their sexual organs, in particular, points to the existence
of a relationship between them ; on the other hand, their asexual
generations exhibit the greatest disagreement, so that it would not
seem admissible to regard the asexual generation of the Pteridophytes
as directly derived from the sporogonium of the Mosses. The
asexual and sexual generations of the Archegoniatae may, however,
be regarded as originally homologous.
The existing Pteridophyta are classified as follows :

1. Filicinae. Ferns. Stem simple or branched, with well-developed,

alternate, often deeply divided or compound leaves. Sporangia either
on the under side of the sporophylls, united in sori or free, or enclosed
in special segments of the leaves.

Sub-Class 1. Filicinae eusporangiatae. Ripe sporangia with firm

wall composed of several layers of cells. Homosporous.
Sub-Class 2. Filicinae leptosporangiatae. Ripe sporangia with
walls one layer thick.

Order 1. Filices. Ferns, in the narrower sense. Homosporous.

Order 2. Hydropterideae. Water-Ferns. Heterosporous. \

2. Equisetinae. Horse-tails. Stem simple or verticillately branched,

with whorled, scale-like leaves forming a united sheath at each node.
Sporophylls peltate, bearing the sporangia on the under side, and
aggregated into a cone at the apex of each fertile shoot.

Order 1. Equisetaceae. Horse-tails. Homosporous.

3. Lycopodinae. Club Mosses. Stems elongated, dichotomously

branched, either forked or forming a sympodium, with leaves, in
many cases greatly reduced, or shortened and tuberous with awl-
shaped Sporangia arising singly in the form of firm-walled
leaves.

capsules either from the stem in the leaf-axils, or from the leaf-base.
Tapetal cells persistent.

Order 1. Lycopodiaceae. Club Mosses. Homosporous.

Order 2. Selaginellaceae. Heterosporous.
Order 3. Isoetaceae. Heterosporous.

There are also various fossil groups, some of which are included
in the above divisions, while some form independent classes.
 CRYPTOGAMS 443

CLASS I.

108
Filicinae. Ferns ( )

The great majority of existing Pteridophytes belong to the

Ferns, taking the group in a wide sense. Two sub-classes are
distinguished according to the structure of the sporangia. The
Eusporangiate Ferns are characterised by sporangia, the thick wall
of which consists of a number of layers of cells ; the sporangium

originates from a group of epidermal cells together with the cells

lying just below the epidermis. The Leptosporangiate Ferns on the
other hand have sporangia which when mature have their wall
formed of one layer of cells ; each sporangium arises from a single
epidermal cell. Stipules, which are found at the base of the frond
in the former group, are wanting in the Leptosporangiatae. Differ-
ences also exist in the prothallus and in the structure of the sexual
organs.

While in earlier geological periods the Eusporangiatae were abundantly

represented, they now include only two families, each with a few genera. They
appear to represent the more ancient type of Ferns and to stand nearest to the
forms from which both Filicinae and Lycopodinae have been derived. The
Leptosporangiatae, from which the Hydropterideae have branched off as a small
group of aquatic or marsh-growing Ferns, may be derived from the Eusporangiatae.
In the Hydropterideae only among Ferns the spores are differentiated into micro-
spores and macrospores.

Sub-Class I. Eusporangiatae.
109
Order 1. Ophioglossaceae ( ).

This family may be placed first since it appears to contain the existing forms
which stand nearest to the primitive ferns. European examples are afforded by
Opltiocjlossum vulgatum, Adder's Tongue (Fig. 403 B) and Botrychium, Moonwort
(Fig. 403 A}. Both have a short stem, from which only a single leaf unfolds
each year. The leaves in both cases are provided with leaf-sheaths, and peculiarly
divided into fertile and sterile segments. In Ophioglossum the sterile leaf-like
segment is tongue-shaped, the fertile segment narrow and cylindrical, bearing
the sporangia in two rows sunk in the tissue. The sterile portion of the leaf
of Botrychium is pinnate, while the fertile segment is pinnately branched, and
thickly beset on the inner side with large nearly spherical sporangia.
Our knowledge of the peculiar monoecious prothalli of the Ophioglossaceae is
largely due to BRUCHMANN ; they are long lived, subterranean, sapropliytic,
-

tuberous bodies without chlorophyll but inhabited by a mycorhizal fungus. In

Ophioglossum (Fig. 403 C) they are cylindrical and radially symmetrical, simple
or branched in Botrychium they are oval or heart-shaped and dorsiventral.
;

The antheridia (Fig. 404) and archegonia (Fig. 405) are sunk in the tissue of
444 BOTANY

the prothallus. The antheridium encloses a large spherical mass of sper-

matozoid mother-cells which are set free when mature by the swelling of the
contents and the breaking down of one of the central cells of the wall. The
spermatozoids have a spirally wound body and numerous cilia a small vesicle ;

is adherent to the spermatozoid (Fig. 404 JE). The antheridia originate from
single superficial cells (Fig. 404 A C), as do also the archegonia (Fig. 405
-

ad

B C
FIG. 403. A, Jiotrychium lunaria. Sporophyte. (J nat. size.) S, Ophwglossum vulgatum. Sporo-
phyte showing the bud for the succeeding year, (i nat. size.) C, Ophioglossum vulgatum.
Prothallus. an, antheridia; ar, archegonia; k, ,young plant with the first root; ad, ad-
ventitious branch ; h, fungal hyphae. (x 15 after BRUCHMANX.)
;

A-G). The slightly projecting neck of the latter opens after the neck-canal-cell
has swollen and disintegrated the oosphere (o) remains in the sunken venter.
;

In many species the embryo leads a subterranean existence for several years.
The primary root is first formed and soon projects from the archegonium (Fig.
403 C, K) later the first leaf and the apical cell of the stem are differentiated.
;

In some species of Botrychium the embryo forms an elongated multicellular

no
suspensor at the end of which the proper embryonic mass is formed ( ). In
this an agreement with the Lycopodinae is evident (cf. Fig. 428).
 CRYPTOGAMS 445

Order 2. Marattiaceae.

This order includes a number of stately tropical Ferns with thickened tuberous
stems and usually very large leaves provided with two stipules at the base.
The sporangia are situated in groups (sori) on the under surface of the leaves,
and are either free (Angiopteris), or united to form an oval capsule-like body, the
chambers of which are the sporangia. The prothallium in contrast to that of the

FIG. 404. Ophioglossum mdgatum. A-C, Stages

in the developmeut of the antheridium from Fin. 405. Ophioglossum vitlgatum. A-C, De-
a superficial cell the upper cell in C gives
; velopment of archegonium D, mature opened ;

rise to the cover -cells, the lower to the archegonium with two .spermatozoids (s) in
mother - cells of the spermatozoids. D, front of the opening h, neck-cells hk, neck-
; ;

Antheridium not yet opened ; d, cover-ci-lls. canal-cells ; o, egg-cell ; 6, basal cell. (After
A'. SpiTiiwto/oids. (After BRUCHMAJTN.) BRUCHMANN.)

Ophioglossaceae is a green, heart-shaped thallus, resembling that of a Liverwort

and growing on the surface of the soil. It is sometimes dichotomously branched.
The sexual organs resemble those of the preceding order but are developed on the
lower surface of the prothallus.

Sub-Class II. Leptosporanglatae.

Order 1. Filices.

The Filices, or Ferns, in the narrower sense of the word, comprise a large
number of genera with numerous species, being widely distributed in all parts of
the world. They attain their highest development in the tropics. The Tree-
Feriis (Cyathea, Alsophila, etc.), which include the largest representatives of
the order, occur in tropical countries, and characterise the special family of the
Cyatheaceae. The stem of a Tree-Fern (Fig. 406) is woody and unbranched it ;

bears at the apex a rosette of pinuately compound leaves or fronds, which are
produced in succession from the terminal bud, and leave, when dead, a. large
leaf scar on the trunk. The stem, resembling that of a Palm in habit, is
attached to the soil by means of numerous adventitious roots. The majority
of Ferns, however, are herbaceous, and possess a creeping rhizome, terminating
usually iu a rosette of pinnate or deeply divided leaves. Such a habit and
growth are illustrated by the common Fern Aspidium filix mas, the rhizome
of which is official (Fig. 407). When young, the leaves of this Fern are
coiled at the tips (Fig. 407, 1, a), a peculiarity common to the Ferns as a
446 BOTANY

whole, and to the Water-Ferns. Unlike the leaves of Phanerogams, those of

the Ferns continue to grow at the apex until their full size is attained. The
leaves of the common Polypodium vulgare are pinnate, and spring singly from
the upper side of the branched rhizome, which creeps amongst Moss or on rocks.
In other cases the leaves may be simple and undivided, as in the Hart's-Tongue

FKJ. 406. Alsophila crinita. A Tree-Fern growing in Ceylon. (Reduced.)

Fern, Scolopendrium vulgare (Fig. 408). In the tropics many herbaceous Ferns
grow as epiphytes on forest trees.
Peculiar brownish scales (paleae, ramenta), often fringed and consisting of a
single layer of cells, invest the stems, petioles, and sometimes also the leaves of
most' Ferns. These are morphologically trichomes.
The sporangia are generally produced in large numbers, on the under side of
the leaves. The sporophylls, as a rule, resemble the sterile, foliage leaves. In a
few genera a pronounced heterophylly is exhibited thus, in the Ostrich Fern,
:

Struthiopteris gennanica (Onoclea- Struthiopteris), the dark brown sporophylls are

 CRYPTOGAMS 447

FIG. 407. Aspidium Jttix mas. ( nat. size). A, Sorus in vertical section (x 20, after KNV) ;

B, Pinna with young sori still covered by the indusia'

1

C, somewhat older sori with withered

indusia. (Slightly magnified.) OFFICIAL.

smaller and less profusely branched, standing in groups in the centre of a rosette
of large foliage leaves.
448 BOTANY

In tlie different families, differences in the mode of development, as well as in

the form, position, and structure of the SPOI:AM;I.\ are manifested.
The sporangia of the Polypodiaceae, in which family
the most familiar and largest number of species are com-
prised, are united in groups or SOKI on the under side
of the leaves. are borne on a cushion-like pro-
They
jection of tissue termed the UKCKPTACLK (Fig. 407 A],
and in many species are covered by a protective
membrane, the IXDUSIUM, which is an overgrowth of
the tissue of the leaf (Fig. 407 B, C). Each spor-
angium arises by the division of a single epidermal
and consists, when ripe (Fig. 409), of a capsule
cell,
attached to the receptacle by a slender multicellular
stalk, containing a largenumber of spores. The wall
of the capsule is formed of a single layer of cells.' A
row of with strongly thickened radial and inner
cells

walls, extending from the stalk over the dorsal side and
top to the middle of the ventral side of the capsule, are
specially developed as a ring or ANNULUS, by means of
which the dehiscence of the sporangium is effected.
This type of annulus is characteristic of the Poly-

podiaceae.
On drying of the wall of the sporangium the co-
hesion of the remaining water in the cells of the
annulus draws in the thin outer walls of these cells ;
this causes the annulus to shorten and determines the
dehiscence of the sporangium by a transverse slit
between the broad terminal cells of the annulus.
When the pull exerted by the cohesive power of the
FIG. 408. water gives way, the annulus returns by its own
Scolopendrium vul-
gare. (J nat. size.) thus effecting the
elasticity to its original position,
dispersal of the spores. The sporangium remains
open owing to the drying and contraction of the thin cell walls.

FIG. 409. Sporangia. A, Aspidium Jttix mas there is a glandular hair at the base ; li and
;
<
,

Alsophila armata, seen from the two sides ; D, Aneimia caudata; E, Osmunda rrgalls.
(A-D x. 70 orig. ; E x 40, after LURSSEN.)

The form and insertion of the sori, the shape of the indusium when present,
or its absence, all constitute important criteria for distinguishing the different
SECT. I CRYPTOGAMS 449

genera. The sori of Scolopendrium are linear, and covered with a lip-shaped
indusium consisting of one cell-layer They are so disposed in pairs, on different
sides of every two successive nerves, that they appear to have a double indusium

opening in the middle. In the genus Aspidium,

on the other hand, each sorus is orbicular in form
and covered by a peltate or reniform indusium
attached to the apex of the placenta a glandular ;

hair is frequently present on the stalk of the

sporangium. The sori of Polypodium vulgare are
also orbicular, but they have no indusia. In the
common Bracken, Pteris aquilina, the sporangia
form a continuous line along the entire margin
of the leaf, which folds over and covers them
with a false indusium.
Besides the Polypodiaceae the Ferns include x

other families, mainly represented in the tropics,

the sporangia of which differ in the construction
of the annulus and in the mechanism of their
11J
dehiscence ( ). The sporangia of the Cyatheacae,
to which family belong principally the tree-like
Ferns, are characterised by a complete annulus
extending obliquely over the apex of the capsule
(Fig. 409 B, 0). The Hymenophyllaceae, often
growing as epiphytes on Tree-Ferns, have also
sporangia, with a complete, oblique, or horizontal i. 410. Trichomanes rigidum. Por-
annulus. The sporangia of the Schizaeaceae, on tion of a prothallus with an arche-

theother hand, have an apical annulus ( Fig. 409 Z>), yoniophore (-4) to which a young
plant is still attached. (After
while in the Osmundaceae, of which the Royal
GOEBEL.)
Fern, Osmunda regalis, is a familiar example,
the annulus is represented merely by a
group of thick-walled cells just below the
apex of the sporangium (Fig. 409 E).
All the
members of the Filices are homospor-
ous. The PROTHALLIUM has usually the form
of a flat, heart-shaped thallus (Fig. 399), bearing
the antheridia and archegonia on the under side.
In certain Hymenophyllaceae (Trichomanes) the
prothallium is filiformand branched, resembling
in structure the protonema of the Mosses, and
producing the antheridia and archegonia on
lateral branches (Fig. 410).
The ANTHERIUIA and ARCHEGONIA (
112
) are
similarly constructed in nearly all Leptosporangi-
atae, and present differences from those of the
Eusporangiate Ferns ; those of Polypodium
Fiti. 411. Polypodium A, vulgare. vulgare (Figs. 411, 412) may serve as a type.
Mature, B, discharged antheridium ;
The antheridia are spherical projecting bodies
1 and 2, ring-
p, prothallium cell ;
(Fig. 411 A, p), arising on young prothallia by
shaped cells 3, lid-cell
;
/>. ;
' .

the septation and further division of papilla-like

sperinatozoids. (A, B x 240 ; C,
D x 540.) protrusions from single superficial cells. When
mature, each antheridium consists of a central
cellular cavity, filled with spermatozoid mother-cells, and enclosed by a wall formed
2 G
450 BOTANY

of two ring-shaped cells (A, 1, 2) and a lid-cell (3). The spermatozoid mother-cells
are produced by the division of the central cell. are discharged from the They
antheridium by the pressure exerted by the swollen ring-cells, and the consequent
rupturing of the lid-cell. Each mother-cell thus ejected liberates a spirally coiled
spermatozoid. The anterior extremity of the spermatozoid is beset with numerous
cilia, while attached to its posterior end is a small vesicle which contains a number
of granules, and represents the unused remnant of the contents of the mother-
cell (Fig. 411 D, C; Fig. 98 ).

The archegonia arise from the many-layered median portion of older prothallia.
They are developed from a single superficial cell, and consist of a ventral portion,
embedded in the prothallium, and a neck portion. The neck, which projects
above the surface of the prothallium, consists of a wall composed of a single layer
of cells made up of four cell rows (Fig. 412 A. B) it encloses the elongated neck- ;

Fi<;. 412. 7'oZt/;>(/iiu vtilyan. -I, Younn iuvhi-pmiuin not vet <>]><MI ; A", nrck-cunal-crll ;

K", ventral canal-cell ; o, egn-cell ; B, mature arche^onium, open, (x '240.)

canal-cell. The ventral portion contains the large egg-cell and the ventral-canal-
cell immediately above it. As the archegonium matures, the canal-cells become
disorganised, and fill the canal with a strongly refractive mucilaginous substance.
This swells on the admission of water, and, rupturing the neck at the apex, is
discharged from the archegonium, which is now ready for fertilisation. The
development of the embryo is represented in Fig. 400.

In certain ferns the sporophyte may originate on the prothallus by a process of

budding or direct vegetative growth the sexual organs are not formed or take
;

no part in the production of the plant (apogamy). Conversely the prothallus may
arise directly, without the intervention of spores, from the tissues of the leaf
ns
(apospory) ( ).

OFFICIAL.Aspidium filix mas, Fiux MAS.

The long brown hairs from the base of the leaf-stalks of various Tree-
silky
Ferns, especially Cibotium Barometz, and other species of this genus, in the East
Indies and the Pacific Islands, are used as a styptic (Penawar, Djambi, Pulu).

Order 2. Hydropterideae (Water- Ferns)

The Water-Ferns include only a few genera, which are more or less aquatic in
habit, growing either in water or marshy places. They are all heterosporous.
 CRYPTOGAMS 451

The macro- and microsporangia do not develop, like those of the Filices, on the
under side of the leaves, but are enclosed in special receptacles at their base,
constituting sporangial fructifications or sporocarps. The wall of the sporangium,
which consists of a single layer of cells, has no annulus.
The Water-Ferns are divided into two families Marsiliaccae, including three
:

genera, and Salviniaceac, with two genera.

Marsiliaceae. To this family belongs the genus Marsilia, comprising about
fifty species, of which M. qiiadrifoliata (Fig. 413) may be taken as an example.
This species grows in marshy meadows, and has a slender, creeping, branched axis,
bearing at intervals single leaves. Each leaf has a long erect petiole, surmounted
by a compound lamina composed of two pairs of leaflets inserted in close proximity.

Kl<:. 413. Miirsili" iiucilrifuliatn. <',

Fn;. 414. I'iltdariu gloltidifera . .-, .Sjxjro-

Young leaf; .-. sponK-irps. (Att<-i ;carp. (After BISCHOFF, reduced.)

BISCHOFK, reduced.)

The stalked oval s]x>rocarps (s) are formed in pairs above the base of the leaf-stalk,
or in other species they are more numerous. Each of them corresponds in develop-
ment to the quadripinnate sterile lamina, but is not divided into pinnae. The
sori within the capsule, disposed in two rows in
of sporangia are enclosed
correspondingly arranged cavities in the young fruit each chamber opens outwards
;

on the ventral side by means of a narrow canal, which eventually becomes closed.
The sporangia are developed originally, as in other ferns, from superficial cells, but
become arched over by the surrounding tissue, and thus subsequently appear as
if formed in internal chambers. As Fig. 413 shows, the young leaves, the develop-
ment of which is as in the Filices, are circinate.
Pilularia also grows in bogs and marshes. It differs from Jfursilia in its
452 BOTANY PART II

simple linear leaves, at the base of which occur the spherical sporocarps, which
uri.se singly from the base of each sterile leaf-segment (Fig. 1 1 1 1.

Salviniaceae. This family contains only free-floating, aquatic plants belonging

to the two genera Salvinia and Azolla. In Salvinia natans, as representative of
the genus, the sparingly branched stem gives rise to three leaves at each
first

node. The two upper leaves of each whorl are oval in shape, and developed as
floating foliage leaves the third, on the other hand, is submerged, and consists of
;

a number of pendent, filamentous segments which are densely covered with hairs,
and assume the functions, of the missing roots. The sporocarps have an entirely
differentmode of development from those of the Marsiliaceae they are spherical, ;

and are borne in small groups on the submerged leaves at the base of the fila-
mentous segments (Fig. 415 A). The sporangia are produced within the sporocarp
from a column-like receptacle, which corresponds in origin to a modified leaf-
segment. The envelope of the sporocarp is equivalent to an indusium it arises :

FIG. 415. Salvinia natans. A, Seen from the side; JB, from above (after HISCHOKK, inluciii) ; f,
an embryonic plant ; msp, macrospore ; )>, prothallium a, stem
; ; In, I/.,, !>.., tin- liist tlirci-

leaves ; bj, the so-called sctitiform leaf. (After PRIN(;SHEIM, x 15.)

as a new growth in the form of an annular wall, which is at first cup-shaped, but
ultimately closes over the receptacle and its sorus of sporangia.
The second genus, Azolla, is chiefly tropical, represented by small floating
plants, profusely branched, and beset with two-ranked closely crowded leaves.
Each leaf consists of two lobes, of which the upper floats on the surface of the
water, while the lower is submerged, and assists in the absorption of water. A
small cavity enclosed within the upper lobe, with a narrow orifice opening outwards,
is always inhabited by filaments of 'an Alga (Anabaena). From the fact that hairs
grow out of the walls of the cavity between the algal filaments, the existence of a
symbiotic relation between the two plants would seem to be indicated. Azolla,
unlike Salvinia, possesses true roots developed from the under side of the stem.
The sporocarps are nearly spherical, and produced usually in pairs on the under
side of the first leaf of some of the lateral branches.
In the structure of the sporangia and spores, and in the development of the pro-
thallia, the Hydropterideae differ in many respects from the Filices. These differ-
ences may be best understood on reference to Salvinia natans ( 114 ) as an
example.
The sporocarps contain either numerous microsporaugia or a smaller number of
SECT. 1 CRYPTOGAMS 453

macrosporangia (Fig. 416 A, ma, mi). In structure both forms of sporangia

resemble the sporangia of the Leptosporangiate Ferns they are stalked, and have,
;

when mature, a thin wall of one cell-layer but no annulus (B, D). The MICRO-
SPORANGIA, enclose a large number of microspores, which, as a result of their
development in tetrads from the mother-cells, are disposed in groups of four ((7),
and embedded in a hardened frothy mass filling the cavity of the sporangium.
This frothy interstitial substance is derived from the tapetal cells, which gradually
lose their individuality and wander in between the spore mother-cells.
The microspores germinate within the microsporaugium, which does not open ;

each germinating microspore puts out a short tubular male prothallium, which
pierces the sporangial wall. The antheridium is developed in this by successive
divisions (Fig. 417). Each antheridium produces four spermatozoids, which are
set free by the rupture of the cell-walls. Although the whole male prothallium is

Fl<;. 416. Salvinic natmis. A, Thrte spoim-arps in median, longitudinal section ; mu, macro-
sporocarp mi, microsporocarp (x 8) ; B, a microsporanginm (x 55) ; C,'portion of the contents
;

of a microeporsnglmn, showing four microspores embedded in the frothy interstitial substance

(x 250); D, a inacrosjKM-aiigiiini and macrospore in median longitudinal section (x 55).

thus greatly reduced, it nevertheless exhibits in its structure a resemblance

to the prothallia of the Filices.
Thf MAruosi'OHANuiA are larger than the microsporangia, but their walls
consist similarly of one cell-layer (Fig. 416 D).Each macrosporangium produces
only a single large macrospore, which develops at the expense of the numerous
spores originally formed. The macrospore is densely filled with large angular
proteid grains, oil globules, and starch grains at its apex the protoplasm is
;

denser and contains the nucleus the membrane of the spore is covered by a dense
;

brown exinium, which in turn is enclosed in a thick frothy envelope, the perinium,
investing the whole spore and corresponding to the interstitial substance of the
microspores, and also formed from the dissolution of the tapetal cells. The
macrospore remains within the sporangium, which is eventually set free from the
mother plant and floats on the surface of the water. On the germination of the
macrospore, a small-celled female prothallium is formed by the division of the
denser protoplasm at the apex, while the large underlying cell does not take part
in the division, but from its reserve material provides the developing prothallium
with nourishment. The sport- wall splits into three valves, the sporangial wall is
ruptured, and the green prothallium protrudes as a small saddle-shaped body.
On it three archegonia are produced, but only the fertilised egg-cell of one of them
 454 BOTANY

a time sunk in the venter of

develops into an embryo l whose foot, remaining for
the archegonium, finally ruptures it (Fig. 418). The first leaf of the germ plant is
shield-shaped (Fig. 415 0).
The development of Azotta ( 11S ) proceeds in a similar manner, but the sporangia
and spores exhibit a number of distinctive peculiarities. The numerous spores of
the microsporangia are aggregated into several nearly spherical balls or massulae,
formed from the interstitial substance derived from the protoplasm ot the
cells. Each massula, enclosing a number of spores, is beset externally with

FlG. 417. Sttlrli'in ?;/.>. Development

of the male prothallium. A, Division
of thei microspore into three cells
/-/// ( x SCO) ; B, lateral view ; r. v.-n-
tral view ofi (mature prothallium '(<
640). Cell / has divided into the pro-
thallium cells n, and p cell //, into the
;

sterile cells l>, c, and the two cells' .j.

each of which has (bnBedltwotsfwnna-
tozoid mother-cells; cell ///, into the 1
Kin. 41s. xf !'< IK i"itnn*. Embryo in longitudinal sec-
sterile cells d, e, and the two cells .-._,. tion ; pr, prothallium ; X, spore-cell ; e, exinium 71,

Tlie cells jt
t sj
ami 83*2 represent two perinium ; spw, sporangial wall ; ar, archegonium ;

antheridia ;
the cells ft, c, :</. e their tmbr, embryo ; /, foot ; MI, bl%, big. the lirst three
wall-cells. (After BELA.IEKF.) leaves : st. ap-x of stem. (After PRINCSHFIM. X 100.)

hook-like outgrowths of the interstitial substance (glochidia). On the rupture

of the sporangia the massulse are set free in the water, and are carried to the

macrospores, to which they become attached. A sporocarp contains one macro-

sporangium, in which only a single macrospore comes to maturity in the course ;

of its development it supplants all the other sporogenous cells, and finally the
sporangial wall itself becomes flattened against the inner wall of the sporocarp,
frequently undergoing at the same time partial dissolution. The macrospore is
enveloped by a spongy perinium whose outer surface exhibits numerous depressions
and protuberances prolonged into filainents. At the apex of the spore the perinium
expands into three pear-shaped appendages, while the upper part of the ruptured
8KCT. I CRYPTOGAMS 455

sporangium remains attached to the spore in the form of an umbrella-like expansion.

The formation of the prothallia is effected in essentially the same way as in

Salvinia, except that only one autheridium with eight spermatozoids arises on
each of the small male prothallia protruding from a massula.
The sporocarps of the Marsiliaceae have a more complicated structure : those of
Pilularia globulifera are divided into four chambers, each with a single sorus in ;

Marsilia they enclose numerous sori (14-18) disposed in two rows. The sori in
both genera contain both micro- and macrosporangia, while those of the Salvini-
aceae always consist of only one kind of sporangium.
In the case of the Marsiliaceae the prothallia are even more reduced, but
otherwise their mode of development is very similar. Each of the minute female
prothallia formed at the apices of the macrospores produces a single archegonium.

CLASS II

116
Equisetinae (Horse-tails) ( )

The Equisetinae, which form an entirely independent class, include only the
one genus Equisetum, comprising 20 species, found widely distributed over the
whole world. Developed partly as land, partly as swamp plants, they may always
be distinguished by the characteristic
structure and habit of the asexual genera-
tion. They have a branching, underground
rhizome on which arise erect, aerial haulms,
usually of annual growth. The rhizome of
the common Horse-tail, Equisetum arvense,
develops also short tuber like branches
-

which function as reservoirs of reserve

material and hibernating organs (Fig. 420).
The aerial haulms remain either simple,
or they give rise to branch whorls, and
these in turn to whorls of a higher order.
All the axes are formed of elongated inter-
nodes they have a central pith-cavity and
;

a peripheral series of smaller air channels.

.The collateral vascular bundles form a
single circle, as seen in transverse section
Fu;. 419. Equisetum arvense. Transverse sec-
(Fig. 419). tion through the stem, m, lysigenic medul-
The leaves of the Equisetiuae, both in
lary cavity e, endodermis d, carinal canals
; ;

their structure and in the manner of their in the collateral bundles vl, vallecular
;

arrangement on the stem, are also char- cavities; hp, sclerenchymatous strands in
the furrows .and ridges eh, tissue of the
acteristically developed. At each node ;

primary cortex containing chlorophyll ; st,

is borne a whorl of scale leaves pointed at
rows of stomata. ( x 11.)
the tips, and united below into a sheath
closely enveloping the base of the internode. The leaves of the successive whorls
alternate regularly with each other. The lateral branches are developed in the
but not having space to grow upwards they pierce the
axils of the scale leaves,
narrow sheath. As a result of the reduction of the leaf laminae, the haulms them-
selves assume the function of assimilation, and for that purpose their cortical
tissue under the epidermis is provided with chlorophyll.
The SPOKANGIA are borne on specially shaped leaves or sporophylls. Like the
456 BOTANY PART II

scale leaves the sporophylh are developed in whorls, but are closely aggregated at

\
Fio. 420. Kquisetum arvense. A, Fertile shoot, springing from the rhizome, which also bears tubers ;
the vegetative shoots have not yet unfolded. F, Sterile vegetative shoot. B, C, Sporophylls
bearing sporangia, which in C have opened. D, Sjx>re showing the two spiral bands of the
perinium. E, Dry spores showing the expanded spiral hands. (A, F, i nat. size. /;. r, 7), K,

enlarged.)

the tips of the erect fertile shoots into a cone (Fig. 420), which is .sometimes
spoken of as a flower, from the correspondence in its structure to the male flower
SECT. I CRYPTOGAMS 457

of the Conifers. The lowest whorl

is sterile, and forms a collar-like protuberance.

The sporophylls C) are stalked and have a peltate expansion, on

(Fig. 420 B,
the under side of which are borne the sac-like sporangia (5-10). In the young
sporangium the sporogenous tissue is surrounded by a wall consisting of several
cell layers, but eventually the tapetal cells of the inner layer become disorganised,
and their protoplasm penetrates between the developing spores. At maturity
the wall of the sporangium consists only of the outermost of the original layers,
the cells of which are provided with annular and spiral thickenings the sporangia ;

thus resemble tl> homologous pollen-sacs of Phanerogams. The dehiscence is deter-

mined by the cohesive force of the diminishing amount of water in the cells of the

II
FJI:. 4'J1. Xijuisetttui pratensf. Female prothallium from the under surface, showing the arche-
I,

k'onia (.4). II, Male prothallium with antheridia (A); d, cover cells of antheridia. (/ x 17,
// x 12. After GOEBEL.)

outer layer and the contraction of the thin parts of the cell walls on drying ( ). m
The sporangia split longitudinally, and set free a large number of green spores,
which are nearly spherical in shape, and have peculiarly constructed walls. In
addition to the intine and exine, the spores are overlaid with a perinium formed
from the protoplasm of the tapetal cells, and consisting of two spiral bands
which are attached to the spores only at their point of intersection (Fig. 420 D).
On drying, the spiral bands loosen and become uncoiled when moistened they ;

close again around the spore. By means of their hygroscopic movements they
serve to hook together the spores, and in this way assure the close proximity
of the unisexual prothallia which the latter produce (Fig. 420 E).
In certain species some of the aerial haulms always remain sterile, branching
profusely, while others which produce the terminal cones either do not branch at
all, or only at a later stage, and then sparingly. This distinction between the
sterile and fertile haulms is most marked in Equisetum arvense and Equisetum

Telmateja, in both of which the fertile shoots are entirely unbranched, and
458 BOTANY PART n

terminate in a single cone (Fig. 420). Resembling in their mode of life a parasite
upon the rhizome, they are otherwise distinguished from the vegetative haulms by
their lack of chlorophyll and their light yellow colour.
Equisetum giganteum, growing in South America, is the tallest species of
the genus ;
its branched haulms, supported by neighbouring plants, attain a

height of over twelve metres, and are about two cm. in diameter.
The spores are all of one kind, and on germination give rise to thalloid
ruuniAM.iA, which are generally dicecious (B'ig. 421). The female prothallia are
larger than the male, and, branching profusely, are prolonged into erect, ruffled
lobes at whose base the archegonia are produced. In structure the archegonia
resemble those of the Ferns, but the upper cells of the four longitudinal rows of
cells constituting the n<;ck arc more elongated and, on opening, curve strongly
outwards. The first leaves of the embryo are arranged in a whorl and encircle the
apex of the stem. The growth of the embryo is effected by the division of a three-
sided apical cell (Figs. 164, 165).
The outer epidermal walls of the stem are more or less strongly impregnated
with silica. In Equisetum hiemale, and to a less degree in Equisetum arvense, the
silicification of the external walls is carried to such an extent that they are used
for scouringmetal utensils and for polishing wood.
Poisonous substances are formed in some species of Equisetum, and hay with
which the shoots are mixed is injurious to cattle.

CLASS III

Lycopodinae (Club Mosses)

Tothe Lycopodinae belong, as their most important and widely
distributed genera, Lycopodium, Selaginella, and Isoetes. They are
distinguished from the other Pteridophyta, of which they resemble
most nearly the Eusporangiate Filices, by their general habit and the
mode of their sporangial development.
and Equisetinae, which
Unlike the fertile leaves of the Filicinae
always bear numerous sporangia, the sporophylls of the Lycopodinae
produce the sporangia singly, at the base of the leaves or in their
axils. Although in many cases scarcely distinguishable from the sterile

leaves, the sporophylls are frequently distinctively shaped, and, like

those of Equisetum, aggregated at the ends of the fertile shoots into
terminal spike-like cones or flowers. Compared with the leaves, the
sporangia are relatively large. They are developed in the same way as
those of the Eusporangiate Filices and Equisetinae, from a projecting
group of cells derived from the epidermis and the underlying tissue ;
while in the other Pteridophyta the sporangia are developed from a
single epidermal cell. The innermost layer of the sporangial wall,
the tapetal layer, is not absorbed. The sporangia have no annulus.
Except in the case of Isoetes, whose spores become free by the decay
of the sporangial wall, they dehisce by longitudinal slits, which divide
the sporangia into two or more valves ; the slits occur where rows of
cells of the wall have remained thin ( ).
m
The sporangia of Lycopodium
 CRYPTOGAMS 459

are homosporous ; those of other Lycopodinae heterosporous. The

heterosporous forms produce only greatly modified and reduced
prothallia ; in the genus Lycopodium, on the other hand, the prothallia
are well developed, and resemble those of the Ophioglossaceae. In the
development of their sexual generation the heterosporous Lycopodinae
resemble in many respects the heterosporous Hydropterideae.
The dichotomous branching of the stems and roots is characteristic
of this class (Figs. 18, 19).

Order 1. Lyeopodiaeeae (
m )

The numerous, widely distributed species of the genus Lycopodium (Club Moss)

FIG. 422. Lycopodium davotiim. A, Old prothallus. B, L'rothallus with .young plant attached.
C, Antheridium in vertical section. D, Spermatozoids. K, Young archegonium, the 'neck
closed.
still F, Open archegoninm ready for fertilisation. G,,Plant bearing cones ( nat. size).
H, Sporophyll with an 'opened sporangium. J, K, Spores from two points of view. (A-F,
after BRUCHMANN.)

most part terrestrial plants in] the tropics epiphytic forms also occur.
are for the ;

In Lycopodium davatum, one of the commonest species, the stem, which is thickly
460 BOTANY PART II

covered with small, awl-shaped leaves, creeps along the ground it branches ;

dichotomously, and gives rise to ascending lateral branches, while from the under
side spring the dichotomously branched roots (Fig. 422). The cone-like flowers, con-
sisting of the closely aggregated sporophylls, are situated in groups of two or more at
the ends of the forked erect shoots. The sporophylls are not like the sterile leaves
in shape ; they are broader and more prolonged at the tip each bears a large ;

reniform sporangium on the upper side

at the base. The sporangium opens
into two valves by a transverse slit,
and numerous minute spores
lets free

(Fig. 422 H). Lycopodium Sclago differs

kj in habit from the other species ; its
G> bifurcately branched stems are all erect,
and the flower-cones are not distinct
from the vegetative region of the fertile
shoots.
The spores of the Lycopodiums are
all of one kind, and in consequence of
their formation in tetrads are of a tetra-
hedral though somewhat rounded shape.
The exine is covered with a reticulate

thickening (Fig. 422 J, K).

The mode of germination and de-
velopment of the sexual generation
have as yet been determined only for
a few species. The prothallia of Lyco-
podium clavatum and the closelyrelated
L. annotinum are small white tuberous
.structures, which live as subterranean
saprophytes. At first top-shaped, they
become converted by the continued
marginal growth into cup-shaped lobed
bodies, which may attain a size of two
centimetres. Long rhizoids spring
from the lower surface, while the upper
surface bears numerous antheridia and
archegonia. In L. complanatum (Fig.
423) the subterranean prothalli are
Fi<;. 423. Lycoixxlium r////>/i''^/H. Prothallus
with antheridia (), archej^onia (or), and a young
turnip-shaped, in L. Selago rounded or
BRUCHMANN. x elongated and cylindrical. The pro-
embryo (k). (After 26.)
thalli of the latter may be developed
on the surface of the soil, in which case they are green. In the case of L,
inundatum, the prothalli of which are found on damp peaty soil, and in the
tropical L. cernuum with erect profusely branched shoots, the prothallia are poor
t

in chlorophyll and are attached to the soil by rhizoids they have the form of ;

small, half-buried, cushion-like masses of tissue, which give rise to green aerial
thalloid lobes. The archegonia occur at the base of these lobes, the antheridia
also on their surface.
The prothallia are all monoecious. The antheridia are somewhat sunk in the
tissue (Fig. 422 C) and enclose numerous spermatozoid mother- cells, in which
small biciliate spermatozoids are formed. The archegonia (Fig. 422 E, F) are
 CRYPTOGAMS 461

constructed like those of the Ferns, but have a shorter neck, whose upper cells
become disorganised on opening. The number of neck-canal-cells differs in the
various species (1, 3, 5, or 6-10).
The embryonic development agrees closely with that of Selayinella (Fig. 428).
A SUSPENSOR is developed it is not, however, laterally inserted, but
; occurs, on
the contrary, at the foot end of the embryo, or between the foot and the apex of
the stem.
The spores of Lycopodium clavatum and other species are sometimes used in
pharmacy.
Order 2. Selaginellaeeae (
11<J
)

To
this order belongs the genus Selaginella, represented by numerous and for
the most part tropical species. They have, as a rule, profusely forked, creeping,
and sympodially branched stems, but occasionally erect, branched stems ;
some

A FIG. 4-2J.Selaginella helvetica. A, macrospor-

line of dehis-
anxium from above showing the
FIG. 424. A, Selaginella. helvetica (from cence (d). B, opened, seen from the side tin- ;

nature, mat. size). B, Selaffinella den- four macrospores, C, have been ejected. /),

ticulata, embryonic plant with macro- microsporangium in the axil of its sporophyll.
spore still attached. (After BISCHOFF, K, the same, opened. F, microspores. (x
magnified.) about 15.)

form moss-like beds of vegetation others, climbing on adjacent plants, possess

;

stems several metres long. In general the Selaginellas are similar in habit to the
Lycopodiums. They have small, scale-like leaves which usually exhibit a dorsi-
ventral arrangement, such as is shown, for example, in the alpine Selaginella
helvetica (Fig. 424), the stem of which bears two rows of small dorsal or upper
leaves, and opposite to them two rows of larger, ventral or under leaves. The
development of a small, membranous ligule at the base of the leaves, on their
dorsal side, is characteristic of the Selaginellas. The rhizophores( 120 ) are organs that
are peculiar to the plants of this order ; they are cylindrical, leafless, shoot-like
structures, which arise exogenously, usually in pairs, from the stem at a bifurcation
462 KOTANY

(cf. p. 50).At their ends a number of endogenous roots are produced, but the
rhizophores are able, when the normal shoots are cut back, to continue their
growth as shoots of ordinary construction. Even below the first leaves of the
seedling plant short rhizopliores are formed, from which the first roots arise
endogenoiisly. In many species of Sclaginella the epidermal assimilatory cells of
the leaves possess, as in Anthoceros, only one large chloroplast ( ). m
As in Lycopodium, the cones or flowers are terminal. Each sporophyll
subtends only one sporangium, which springs from the stem above the leaf-axil.
The same spike bears both macrosporaugia and microsporangia. Each macro-
sporangium (Fig. 425 A-C) contains only four macrospores, which are produced
by the growth and division of a single spore-mother-cell all the other mother- ;

cells originally developed ultimately disappear. On account of the increasing size

Klo. 4-26. A-E, Selugiitdla stolonijera, successive .stages in the ^ermiiiation of a Imicrospore.;
p, prothalliuin-cell ; if, wall-cells of aiitheridium ; six>rmatogenous cells ; A, B, D, lateral,
.-.

C, dorsal view. In E the prothal Hum-cell is not visible, the disorganised wall-cells enclose
the spermatozoid mother-cells 7-".
ipermttOBOid*
; ot'lSel"'.ii it'll" i-n*i>iilutii. (After HELAJEKK.
A-K x 640, F X 780.)-

of the spores. the spherical macrosporaugia become nodular. Opening occurs along
definite lines of dehiscence, the wall splitting into two valves, which curve back
from a boat-shaped basal portion. The spores are ejected by the pressure of the
contracting boat-shaped part and the valves. Numerous spores are formed in the
flattened microsporangia. The mode of dehiscence is similar in these also, but the
boat-shaped portion of the wall is smaller, the valves extending to the base.
The microspores begin their development while still enclosed within the
sporangium. The spore first divides into a small lenticular vegetative cell, which
corresponds to the rhizoid cell of Salvinia, and into a large cell, which divides
successively into eight sterile peripheral cells and two or four central sperma-
togenous cells (Fig. 426 A). By the further division of the central cells numerous
spermatozoid mother-cells are formed (B-D). The peripheral cells then break
down and give rise to a mucilaginous substance, in which is embedded the central
mass of spermatozoid mother-cells (E). The small prothallium-cell, however,
persists. Eventually the wall ruptures, and the mother-cells, thus liberated,
set free the club-shaped biciliate spermatozoids (F). The reduction here exhibited
 CRYPTOGAMS 463

in the formation of the male prothallium resembles that shown by the Hydro-
pterideae.
The macrospores in some species
similarly begin their development
within the sporangia. After the ^=4
division of the nucleus-which lies _ { \^8SKtSM^ICQ}S&^ ar
in the peripheral cytoplasm at the

apex into daughter nuclei and their

distribution in the apical cytoplasm,
the formation of cell walls begins.
In this way, progressing from apex
becomes filled by a
to base, the spore

process of multicellular formation,

with large prothallium-cells. At the
same time, and proceeding in the
same direction, there begins a further
division of these cells into smaller
cells. In some species the apical disc itpm
of tissue is formed first, and is separ-

ated by a thickened wall or diaphragm Kl(

.
4 27 ._ sria ^ Md j tt uartwii. Female prothallium
from the rest of the cavity of the protruding from tlie apex of the ruptured macro-
spore ;
cell-formation occurs in this sj>ore ;spm, wall of macrospore ; ar, an unfertilised
later. In the tissue at the apex, con- arehegonium eiiibi, emb^, two embryos, with sus-
;

pensors et, sunk in the tissue of the prothalliuni.

sisting of small cells, the rudiments (After PFEFFER, modified, x 124.)
of a few archegonia appear, often even
before the formation of the prothalliuni has been completed. The archegonia are
usually not formed until the spores have been discharged from the sporangium.

Fi<;. 4'J*. tieliiyiitellu Murtensii. Longitudinal section of an embryo before its sejiaratiou from
the spore ; et, suspensor : w, root /, toot
; hi. le.-ives
: Hy, lij^ules ft, apex of stem. (After
; :

I'FKKFER, x 165.)

The wall of the spore eventually bursts at the apex, and the
prothallium
becomes partially protruded. The fertilisation of one or two archegonia, which
464 BOTANY

then takes place, is followed directly by the segmentation of the fertilised egg-cells
and the formation of the embryos (Fig. 427).
The embryogeny of Selayinella recalls that of Lycopodium. The egg-cell is
divided by the formation of a transverse wall into two cells the upper and ;

larger cell increases considerably in size, and gives rise, by the division of its
lower portion, to a suspensor (Fig. 428 ct), while the lower cell, by repeated
division, develops into an embryo, provided with two primary leaves and further
.segmented into stem, root, and foot (bl, st, w, f). The foot, in this instance, has
a different position and origin than in Lycopodium. Each primary leaf has, even
formed as an outgrowth of the leaf-base.
at this stage, a ligule (lig)
The suspensor is perpendicular to the axis of the embryo its function is to ;

push the embryo into the tissue of the prothallium, with which the foot, the organ
of absorption, is thus kept in close contact: The stem apex, with the first pair of
leaves, eventually grows upwards, and the root also extends beyond the macro-
spore. As the foot still remains in the prothallium the young plant continues
united to the spore, and presents the appearance of a phanerogamic seedling with
the seed still attached (Fig. 424 B).

1
Order 3. Isoetaeeae ( --)

isolated genus Isoetes must be regarded as a persistent branch of an ancient

The
group of plants. It exhibits some affinities with the eusporangiate Ferns. The
species of Jsoctcs are perennial plants, growing
either on damp soil or submerged in water. The
stem short and tuberous, terminating below
is

in a tuft of dichotomously branching roots,

;.430. A-F, Isoetes setacea (x 640). A, microspore

seen from the side. B-D, segmentation of the spoiv :

p, prothallial cell; w, the- fonr cells of the wall ;

s. .sjii'Miiato.uenous the four spermatozoid

cells. 7-,',

mother-cells ure .surrounded by the disorganised oeilfl

of the wall ;lsurface view. F, the same in side view.
0, Isoetes Malinverniana, spermatozoid. (x 780.
. 4-_".. Isoetes lacustris. (i nat. size.) After BELAJEFF.)

and above in a thick rosette of long, stiff, awl-shaped leaves (Fig. 429). The
leaves are traversed longitudinally by four air-passages, and expand at the base
into a broad sheath. On the inner side of the leaves, above their point of
insertion, is an elongated pit, the fovea, containing a large sessile sporangium.
 CRYPTOGAMS 465

A ligule, in the form of a triangular membrane, is inserted above the fovea.

Isoetes thus differs greatly in habit from the other genera, but resembles Selaginella
in the development of a ligule.
The macros porangia are situated on the outer leaves of the rosette ;
the micro-
sporangia on the inner. Both are traversed by transverse plates of tissue or
trabeculae, and are in this way imperfectly divided into a series of chambers. The
by the decay of the sporangial walls.
spores are set free
The development of the sexual generation is accomplished in the same way as
in Selaginella. The reduced male prothallium (Fig. 430) arises similarly within
the spore, by the formation of a
small, lenticular, vegetative cell,
and a larger cell, the rudiment of a
single antheridium. The larger cell
divides further into four sterile per-
ipheral cells, which completely en-
close two central spermatogenous
cells. From each of the latter arise,
in turn, two spermatozoid mother-
cells, four in all, each of which, when
liberated by the rupture of the spore
wall, gives rise to a single, spirally
coiled, multiciliate
spermatozoid.
The female
prothallium (Fig. 431)
just as in Selaginella, also remains FIG. 431. Isoetes echinospora. A, Female prothallium ;

enclosed within the macrospore, ar archegonium ; o, egg-cell. B, C, development

>

of the archegonium from a.superficial cell A neck-

and is incapable of
independent
;

cells hk, neck canal-cell ; h, ventral canal-cell ;

;

growth. It shows similarly an ap- 0> egg . ce ii. (After CAMPBELL, x 250.)
proach to the Conifers, in that the
nucleus first divides into numerous, parietal daughter-nuclei before the gradual
formation of the cell walls, which takes place from the apex of the spore to the
base. As a result of this process the whole spore becomes filled with a pro-
thallium, at the apex of which the archegonia are developed. The embryo has
no suspensor. In the structure of the embryo and of the spermatozoids Isoetes
differs from the other Lycopodineae, and resembles in some respects the euspor-

angiate Ferns.

123
The Fossil Cryptogams ( )

The remains of cryptogamic plants of former geological periods afford no

evidence as to the phylogenetic relations of the classes of Thallophyta and
Bryophyta. Intermediate forms between Algae and Bryophyta and between the
latter group and the Fteridophyta are, as yet, unknown. On the other hand
Phytopalseontology has made us acquainted with interesting, long-extinct types
of Pteridophytes, which serve to'complete the classification of the existing Ferns,
Horse-tails, and Club-mosses, and in part afford a transition from the Ferns to
the Gymnosperms.
I. The great majority of Thallophytes are, on account of their delicate structure,

not adapted to be preserved in the fossil condition. The absence of remains of

many classes of Thallophytes is therefore no ground for concluding that they did
not exist in earlier periods. Even in the Silurian rocks remains of Algae are
found, though they do not permit of their relationship being determined. The
remains of calcareous Algae belonging to the Siphonales are, owing to their good
2 H
466 BOTANY PART n

preservation, among the best known ; of these, forms are known from the Tertiary
strata back to the Silurian, while the Corallineae, which are calcareous Red Sea-
weeds, appear from the Upper Jurassic onwards. Among unicellular Algae the
Diatomaceae, which have a silicified cell wall, are well preserved. The fossil forms
frequently belong to existing genera, and occur from the Jurassic onwards.
especially in the Cretaceous and Tertiary strata, often forming thick layers
of

Kieselguhr (diatomaceous earth). From Tertiary times onwards Characeae are

abundant, and occasional remains are found as far back as the Muschelkalk. Most
of the existing groups of Algae are only to be distinguished with certainty from
Tertiary strata onwards.
Bacteria must have played their part in causing the decomposition of organic
substances from very early times they can be distinguished in vegetable remains
;

from Carboniferous rocks. The Eumycetes and probably the Myxomycetes also, were
present in Carboniferous times Ascomycetes living on leaves and stems are found
;

from the Carboniferous onwards in all strata. Remains of existing genera of

Lichens appear in the Tertiary rocks.
II. Bryophyta. Most of the examples of this group, which is rarely met with
in the fossil condition, come from the Tertiary strata and resemble existing genera

closely. Only isolated examples of Liverworts and Mosses occur in the older rocks
(Jurassic, Upper Triassic).
III. The Pteridophyta extend back to Silurian times, but were most highly

developed in the Carboniferous period, when they formed the main mass of the
land vegetation. With the advent of Gyninosperms, and later of Angiosperms,
their relative importance in this respect diminished.
1. The class of Filicinae was represented by forms belonging to both sub-

classes at the end of the Silurian period, and large numbers of species occur in
the Carboniferous strata. The organisation of these ancient types is essentially
similar to that of the existing forms. Most of the recent families were represented,
some (e.g. Marattiaceae) more fully than at the present time. The Cycadofilices,
fern-like plants with secondary thickening, which already bore seeds, appear to
have given rise to the Phanerogams, or at least to the Cycadaceae. The other
classes of Pteridophyta do not appear to have given origin to higher forms.
The Hydropterideae are known with certainty from the Tertiary rocks, but
Salvinia and Marsilia can be traced back to the Chalk.
2. The class of the Equisetinae represented at present by the single genus

Equisetum, which can be traced back to the Triassic period, was much better
developed in Palaeozoic times. The large order Calamarieae consisted of plants
resembling the Horse-tails in general habit, but in some cases attaining the size
of trees 30 metres high the hollow stem bore whorls of branches at the nodes,
;

was covered with a periderm, and underwent secondary thickening. The leaves
(Annularia] stood in alternating whorls their form was narrowly lanceolate and
:

at their bases they united into a sheath. In the most ancient type, Archaco-
calamites, they were dichotomously divided. The cones or flowers (Calamost<iclii/s>
had in some the same structure as those of Equisetum in most cases they w ere
;

more complicated, whorls of scale-leaves alternating with the sporophylls. At least

some of the Calamarieae were heterosporous.
3. The Lycopodinae were also
abundantly represented in Palaeozoic times,
especially by the two great extinct
orders, the Sigillarieae and the Lepidodendreae.
The found from the Culm onwards, are most numerous in the Car-
Sigillarieae,
boniferous period, and persist with one species in the Bunter Sandstone. They
were stately trees, with but little branched, pillar-like stems, which grew in thick-
 CRYPTOGAMS 467

ness. They had long narrow leaves, which when they fell off left longitudinal
rows of hexagonal leaf-scars on the surface of the stem. Long-stalked, cone-like
flowers were borne on the stem ;
only one kind of spore was contained in the
sporangia.
The Lepidodendreae extend from the Lower Devonian to the Rothliegende, but
are also best developed in the Carboniferous period. They were tree-like plants
with dichotomously branched stems which grew in thickness. The leaves, which
attained a length of 15 cm., were spirally arranged and seated on rhombic leaf-
cushions. The cone-like flowers (Lepidostrobus) were borne on the ends of branches
or sprang from the stem itself; each sporophyll bore a single sporangium, which
contained either inacrospores or microspores.
Smaller Lycopodiaceae, the predecessors of the existing species of Lycopodium,
were already present in the Carboniferous flora, while Isoetes is only known with
certainty from the lower Cretaceous strata.
4. The small class of the Sphenophyllinae, which existed from the Devonian

to the Permian and then died out, possesses special morphological interest since it

occupies an intermediate position between Lycopodinae and Equisetinae. Possibly

it is the least altered from the common ancestral form of these three groups. This
especially holds for the most ancient type Cheirostrobus, which occurs in the Lower
Carboniferous rocks. The cones of this genus had a very complicated structure,
reminding one of the calamarian cone, while the anatomy exhibits an approxima-
tion to the type of Lepidodendron.
The species of Sphenophyllum were plants with elongated stems, bearing
superposed whorls of obcuneate or more or less dichotomously divided leaves.
The cones were large and terminal, resembling those of Equisetum each sporo-
;

phyll bore two or three homosporous sporangia. These plants have been regarded
as aquatic, but the structure of the elongated thin stem, with a triangular mass of

primary xylem and subsequent secondary thickening, rather suggests that they
were climbing plants which grew on land.
 PART II

SPECIAL BOTANY

SECTION II

PHANEROGAMIA
 SECTION II

PHANEKOGAMIA
The Transition from the Cryptogams to the Phanerogams. The
old names, Cryptogams and Phanerogams, are here retained partly on
historical grounds and also because there are no better and equally
short terms for the two great grades of the vegetable kingdom. The
sharp distinction which formerly appeared to exist between the two
groups has, however, as our knowledge has advanced, become less
marked. The Phanerogams appear as a continuous development
from heterosporous Vascular Cryptogams. The recognition of this
owing, in the first place, to the work of W. HOFMEISTER ( ).
1
is

been seen (p. 422) that the existence of an alternation of

It has
is an essential character, common to both
generations Bryophytes and
Pteridophytes ; the life history exhibits a regular alternation of a
sexual with an asexual generation. While in the Bryophyta only
the sexual generation attains an independent existence, in the
Vascular Cryptogams this stage appears as the inconspicuous pro-
thallium. The asexual generation, on the other hand, which in the
Moss was represented by the sporogonium dependent throughout its
lifeon the sexual plant, becomes physiologically independent in the
Pteridophyta. In them it appears as the conspicuous plant, the Fern
or Horsetail, and bears leaves, on some of which, the sporophylls,
the sporangia develop. A shoot bearing a number of sporophylls,
and frequently with other leaves forming an outer investment, is
known as a FLOWER. Thus Equisetum (Fig. 420, p. 456) affords a
good example of a flower of simple construction, in which the
sporophylls are all alike.
The appearanceof heterospory (p. 441) marks a most important
advance the sexual differentiation, which in homosporous forms did
;

not appear until the sexual generation, is evident in the asexual

plant. The latter produces male sporangia (the microsporangia) and
female macrosporangia. The function of the sexual generation is
limited to the production of the male or female sexual organs, and it
471
472 BOTANY PAKT n

undergoes still further reduction. In the germination of the micro-

spores only a single, vegetative prothallium-cell is to be recognised,
the remainder of the small prothallium representing one or more
antheridia. The female prothallium, which in Salvinia still becomes
green and emerges from the macrospore, in Selaginella and Isoetes has
lost the power of independent nutrition. The prothallium begins
its development while still within the
macrosporangium of the parent
plant, and the macrospore after being set free only opens in order to
allow of the access of the spermatozoids to the archegonia.
Thus within the series of Vascular Cryptogams a reduction
of the sexual generation can easily be traced ; this reduction is
carried further in those Phanerogams which stand nearest to the
2
Cryptogams ( ).
The MACROSPORE, which in the Phanerogams has for long been
termed the EMBRYO-SAC, remains enclosed in the MACROSPORANGIUM

C
Km. 432. A, Atropous B, anatropous
; '', eampylotropous ovules.
;

(Diagrammatic and magnified.)

or OVULE. The latter consists of the NUCELLUS, from the base of which
(the CHALAZA) one or two INTEGUMENTS arise; these grow up as tubular
investments of the nucellus and only leave a small passage, the
MICROPYLE, leading to the tip of the latter. The ovule is attached
to the MACRO-SPOROPHYLL or CARPEL by a stalk or FUNICULUS, which
is often very short. The region to which one or more ovules are
attached is called the PLACENTA. If the nucellus forms the direct
continuation of the funiculus the ovule is termed straight or ATROPOUS.
More frequently the funiculus is sharply curved just below the
chalaza so that the ovule is bent back alongside its stalk (ANATROPOUS
ovule). The line of junction of the funiculus with the outer integu-
ment is still recognisable in the ripe seed and is termed the RAPHE.
Lastly the ovule itself may be curved, in which case it is spoken of as
CAMPYLOTROPOUS. The three types are diagrammatically represented
in Fig. 432 A-C.
As a rule only one embryo-sac is contained in an ovule. In the
same way as the four macrospores originate by the tetrad division in
 PHANEROGAMIA 473

the macrosporangium of Selaginella, in the macrosporangium (ovule) of

the Phanerogams there is usually a single embryo-sac mother-cell
which divides into four daughter-cells ; three of these do not develop
further, while the fourth becomes the embryo-sac. The formation of
the PROTHALLIUM or ENDOSPERM and of the ARCHEGONIA or EGG-CELLS
differs in the several classes of Phanerogams. The fertilised ovum
grows into the embryo while enclosed within the macrospore and
still
at the expense of the parent plant. When the embryo has reached a
certain stage in its development, which is different and characteristic
in different plants, its growth is arrested, and after the separation
from the parent plant it undergoes a period of rest. It is still
surrounded by the other portions of the macrosporangium, viz. the
prothallium or endosperm, the nucellus (if this still persists), and the
seed coat formed from the integuments. THE COMPLETE STRUCTURE

Fio. 433. Part of section through one of the coty-

ledons of the Pea, showing cells with reserve FIG. 434. Cell from the endosperm of
material, am, Starch grains ;al, aleurone grains; Phytelephas tnacrocarpa, with re.srnv
p, protoplasm; n, nucleus, (x 160.) cellulose, (x 225.)

DERIVED FROM THE OVULE IS TERMED A SEED, AND THE FURTHER

DEVELOPMENT OF THE UNOPENED MACROSPORANGIUM TO FORM A SEED
is CHARACTERISTIC OF ALL PHANEROGAMS. As seed-plants or Sperma-
phyta the latter may be contrasted with the seedless Cryptogams.
The several parts of the ripe seed are thus the EMBRYO, the
PROTHALLIUM or ENDOSPERM surrounding it, the remains of the
NUCELLUS which is distinguished from the endosperm as PERISPERM,
and the SEED-COAT. The cells of the endosperm and the perisperm
when present are stored with reserve materials such as starch, proteid
substances, or fatty oils (Figs. 433, 434) ; cellulose in the form of
greatly thickened cell walls may also be a reserve substance. When
the seed after a period of rest commences to germinate, these sub-
stances are utilised by the young plant until it becomes capable of
independent nutrition. Since the nucellus is as a rule completely
obliterated by the enlarging endosperm, few seeds possess a perisperm
(Fig. 435). Frequently no endosperm is present in the seed, and the
storage of reserve material takes place throughout the embryo (Fig.
436) or in its swollen seed-leaves or COTYLEDONS.
474 BOTANY

The SEED-COAT presents great variety in its appearance. It is

always constructed to protect the tender parts within against drying or
injury, and usually shows layers of cells with their walls thickened
and lignified or suberised. On the outside of the seed-coat the scar
of the funiculus can be distinguished, and is known as the HILUM the ;

MICROPYLE can also be seen, and in anatropous seeds the RAPHE mark-
ing the line of union between the funiculus and the integument, and
extending from the hihim to the chalaza. In some cases an ARILLUS is
developed from the funiculus this is a succulent and usually brightly
;

coloured structure, and stands in relation to the dispersal of the seed by

animals. An outgrowth surrounding the micropyle is found in some
A v

Fio. 436. Capsflla burssa pastoris. A, Longi-

tudinal section of a ripe seed; /(, hypocotyl;
FK;. 435. Seed of Hyoscyamus niger, showing
.1.
c, cotyledons ; v, vascular bundle of thr
the dicotyledonous embryo embedded in the funicle(x 26); B, longitudinal section of
endosperm B, seed of Elettaria Cardamoimim,
; the seed-coat after treatment with water ;

enveloped by a thin aril ; the white, mealy t, the swollen epidermis; o, brown, strongly

perisperm next to the seed - coat encloses thickened layer *, compressed layer of
;

an oleaginous endosperm (shaded), in which cells ; a, the single porsisting layer of

the monocqtyledonous embryo lies embedded. endosperm cells containing alem-onc grains
(After BERG and SCHMIDT.) (x 250).

seeds, especially those of the Euphorbiaceae, and is termed a CARUN-

CULA (Figs. 437, 438).
The effect of fertilisation is not only seen in the macrosporangia
but extends to the macrosporophylls or carpels. The structures of
very various form which are formed from the carpels (often together
with the persistent calyx and the floral axis) are called FRUITS, and
serve primarily to protect the developing seeds. The different forms
of fruit will be frequently referred to in the special accounts of the
classes and orders below.
The GERMINATION of the seed, i.e. the resumption of growth at the
end of the resting period, must also be briefly considered. The chief
condition of germination is the absorption of water. The embryo
resumes its growth, which has for a time been arrested, ruptures the
seed-coat and sends its root down into the soil. When it is in this
way firmly fixed in position the cotyledons become free from the
seed and expand in the light when they remain within the seed-
;
 PHANEROGAMIA 47&

coat as in the Pea and Horse-chestnut, the plumule (i.e. the young
shoot) is quickly developed at the expense of the reserve materials
stored in the cotyledons (cf. Fig. 169; pp. 161 and 320).
An account of the male sexual generation of the Phanerogams
must now be given for comparison with the above short sketch of
the development of the female generation.
The MICROSPORES of the Spermaphyta are called POLLEN-GRAINS,
They are formed in large numbers within the MICROSPORANGIA or
POLLEN-SACS, which are borne singly or in numbers on the MICRO-
SPOROPHYLLS or STAMENS. The part of the stamen which bears the
pollen-sacs usually clearly distinguishable and is called the ANTHER.
is

The development of the pollen-sac (Fig. 439) commences with

divisions parallel to the surface taking place in cells of the hypo-

Fiu. 437. .4, Seed of Papaver Khiirits :

h,
the hilum ; B, seed ot'Corydalis ochro-
leuca ; m, micropyle ; c, carnncula : FIG. 438. A, Myristica fragrans, seed from which the
C, seed of Chelidonium majus D, seed
: arillus (ar) is partly detached. B, Myristica argcntea,
of Nymphaea alba with its arillus. seed after removal of the arillus ; ch, chalaza r,
;

(After DUCHARTBE.) raphe: h, hilum. (After WARBUIU;. } nat. si/c.)

dermal layer ; this separates the cells of the primary archesporium

from an outer layer of cells. The latter give rise to three layers of
cells, the outermost of which is the fibrous layer, the innermost the

tapetum, while the intervening layer is later crushed. The arche-

sporium after undergoing a number of divisions forms the pollen-
mother-cells, each of which divides as in Pteridophytes into four
daughter-cells. These are the pollen -grains, and are spherical or
ellipsoidal in shape and provided with a cell wall ; an external
cutinised layer (the EXINE), and an inner cellulose layer, rich in
pectic substances (the INTINE), can be distinguished in the wall.
While the male sexual cells of all archegoniate plants are depend-
ent on water for their conveyance to the female organs, the transport
egg-cells is brought about
of the pollen-grains to the in Seed-plants

by means of the wind or by animals. However far the reduction of

the male prothallium has proceeded and even in the case of the
heterosporous Pteridophyta only single sterile cell was present,
a two
476 BOTANY

constituent parts are always distinguishable in the germinating pollen-

grain ; these are a VEGETATIVK CELL which grows out as the POLLEN-
TUBE, and an ANTHERIDIAL MOTHER-CELL which ultimately gives rise
to two GENERATIVE CELLS. The pollen-tube, the wall of which is

continuous with the intine of the pollen-grain, ruptures the exine

and penetrates, owing to its chemotropic irritability, into the tissue
of the macrosporangium
(cf. p. 286). The antheridial mother-cell
passes the pollen -tube and sooner or later gives rise to two
into
generative cells which reach the embryo-sac and egg-cell by passing

Fio. 439. Hcmerocallis fulva. A, Transverse section of an almost ripe anther, showing the locnli
ruptured in cutting p, partition wall between the loculi a, groove in connective /, vascular
; ; ;

bundle ( x 14) B, transverse section of young anther ( x 28) C, part of transverse section of a
; ;

pollen-sac; pm, pollen -mother- cells t, tapetal layer, later undergoing dissolution
; e inter-
;

mediate parietal layer, becoming ultimately compressed and disorganised /, parietal layer of
;

eventually fibrous cells; e epidermis (x 240); D and E, pollen-mother-cells after division

(x 240).

along the pollen-tube. The name Siphonogams has been applied to

the seed-plants on account of the common character of the group
afforded by the formation of a pollen-tube.
The results reached by the above survey may be summarised by
saying that the Phanerogams continue the series of the Archegoniatae
and agree with the latter in exhibiting an alternation of generations.
While the asexual generation becomes more complex in form and
more highly organised, there is a corresponding reduction of the sexual
generation. The female sexual generation is enclosed throughout its
whole development in the asexual plant, and only becomes separated
from the latter in the seed, which further contains as the embryo the
commencement of the succeeding asexual generation. The exhaustive
investigations made of recent years into the phenomena of the
Sexual Generation Asexual Generation
478 BOTANY FAIIT 11

reduction division in the spore -mother -cells of archegoniates and

Phanerogams have resulted in a confirmation of the limits of the two
2a
generations in the latter ( ). The number of chromosomes charac-
teristic of any plant is diminished to one half during the divisions
that lead to the origin of the sexual generation, and the full number of
chromosomes is not again attained until fertilisation takes place. The
asexual generation has always the double number, the sexual genera-
tion the single number of chromosomes. The constant progression
of the former generation at the cost of the latter might be placed in
relation with this difference between them ; a strict demonstration of
such a connection is, however, not possible.
The Spermaphyta are divided into two classes ( 3 ) which differ in
their whole construction: (1) the Gymnosperms, with naked seeds;

(2) the Angiosperms, with seeds enclosed in an ovary.

The names of these classes indicate the nature of one of the most
important differences between them. THE CARPELS OF THE ANGIO-
SPERMS FORM A CLOSED CAVITY, THE OVARY, WITHIN WHICH THE
OVULES DEVELOP. SUCH AN OVARY IS WANTING IN THE GYMNO-
SPERMS, THE OVULES OF WHICH ARE BORNE FREELY EXPOSED ON
THE MACROSPOROPHYLLS OR CARPELS.
The Gymnosperms are the phylogenetically older group. Their
construction is simpler and in the relations of their sexual generation
they connect directly with the heterosporous Archegoniatae they ;

might indeed be perhaps best treated as belonging to this group.

The Angiosperms exhibit a much wider range in their morpho-
logical and anatomical structure. The course of their life-history
differs considerably from that of the Gymnosperms, and without the
intermediate links supplied by the latter group, the correspondence
with the life-history of the Archegoniatae would not be so clearly
recognisable.
These conclusions are confirmed by the evidence afforded by
Palaeobotany. Gymnosperms them are found
or forms resembling
along with what appear to be intermediate forms between the Gymno-
sperms and the Pteridophyta in the fossiliferous rocks of the Devonian,
Carboniferous, and Permian formations. The Angiosperms are, on
the other hand, first known from the Cretaceous formation.

Table of the Classes and the most Important Orders

f
and Families of Phanerogams
Class I. Gymnospermae
1. Cycadinac Fam. Cycadaeeae.
:

2. Ginkgoinac Fam. Ginkgoaceae.

:

3. Coniferae : Fam. Taxaceae, Pinaceae.

4. Gnetinac : Fam. Gnetaceae.
SECT, ii PHANEROGAMIA 479

Class II. Angiospermae

Sub -Class A. Monocotylae

1. Helobiae : Fain. Alismaceae. Juncaginaceae. Potamogetonaceae. Xaiadaceae.

Hydrocharitaceae.
2. Glumiflorae, : Fam. Cyperaceae. Grainineae.
3. Spadiciflorae Fam. Typhaceae. :
Sparganiaceae. Pandanaceae. Palmae.
Araceae. Lemuaceae.
4. Enantioblastae : Fam. Commelinaceae.
5. Liliiflorae : Fam. Juncaceae. Liliaceae. Amaryllidaceae. Iridaceae. Dio-
scoreaceae. Bromeliaceae.
6. Scitamineae : Fam. Musaceae. Zingiberaceae. Canuaceae. Marantaceae.
7. Gfynandrac : Fam. Orchidaceae.

Sub- Class B. Dicotylae

(a) Choripetalae
1. Piperinae Fam. Piperaceae. -4-
:

2. Juglcmdifiorac Fam. Juglandaceae. Myricaceae. :

3. Saliciftorae Fam. Salicaceae.

:

4. Querciflorae Fam. Betulaceae. Cupuliferae.

:

5. Urticinae Fam. Ulmaceae. Moraceae. Cannabinaceae. Urticaceae.

:

6. Centrospermac Fam. Polygouaceae. Chenopodiaceae. Caryophyllaceae.

:

7. Polycarpicae^? Fam. Nymphaeaceae. Ceratophyllaceae. Ranunculaceat'.

Magnoliaceae. Auonaceae. Myristicaceae. Menispermaceae. Berberidaceae.
Lauraceae.
s. Fam. PajiaveraceaeP Crucifera^". Capparidaeeae. Resedaceae.
Rhocadinac :

9. Fam. Sarraceniaceae. Nepenthaceae. Droseraceae.

Insectivorae :
-

10. Saxifraginae Fam. Crassulaceae. Saxifragaceae. Hamamelidaceae. -/

:

11. Ilosiflorae Fam. Rosaceae. :

C^2. Leguminosae Fam. Mimosaceae. Caesalpiuiaceae. Papilionaceae.

:

13. Gruinales Fam. Geraniaceae. : Oxalidaceae. Linaceae. Balsaminaceae.

Erythroxylaceae. Zygophyllaceae. Rutaceae. Simarubaceae. Burseraceae.
Polygalaceae.
14. Tricoccae : Fam. Euphorbiaceae. Callitrichaceae.
15. Sapindinac Fam. Buxaceae. Anacardiaceae. Aquifoliaceae.
: Celastraceae.
Aceraceae. Sapindaceae.
16. Frangulinae Fam. Rhamuaceae. Vitaceae.
:

17. Columnifcrae Fam. Tiliaceae. Sterculiaceae. : Malvaceae.

18. Cistiftorae : Fam. Ternstroemiaceae. Guttiferae. Dipterocarpaceae. Cistaceae.
Violaceae.
19. Passiftorinae : Fain. Passifloraceae. Caricaceae. Begoniaceae.
20. Opuntinae Fam. Cactaceae.
:

21. Thy melaeinae Fam. Thymelaeaceae. Elaeagnaceae.:

r
. 'I'l.
Mijrtiflorae Fam. Lythraceae. :Melastomaceae. Ouagraceae. Halorrha-
Rhizophoraceae.
gidaceae. Combretaceae. Myrtaceae. Punicaceae.
23. Umbelliftorae Fam. Cornaceae. Araliaceae. Umbellit'erae.
:

24. Hysterophyta Fam. Ari.stolochiaceae. Santalaceae.

: Lorantliaceae. Bal-
anophoraceae. Ratttesiaceae.
 480 BOTANY

(b) Sympetalae
(a) Pcntacyclicae.
1. Ericinae : Fain. Ericaceae. Pirolaceae.
'2.
Diospyrinae : Fam. Sapotaceae. Ebenaceae. Styracaceae.
:5. Primulinae : Faiu. Primulaceae. Plumbaginaceae.
() I'etmcyclicae.
4. GContortae : Fam. Oleaceac. Loganiaceae. Gentianaceae. Apocynaceae.
Asclepiadaceae.
5. Tubifiorae : Fam. Convolvulaceae. Polemoniaceae. Hydrophyllaceae. Bor-
raginaceae. Verbenaceae. Labiatae.
6. Personatae : Fam. Solaiiai-cai-. Scrophulariaceae. Orobanchaceae. I'.ig-

noniaceae. Gesneriaceae. Lentibulariaceae. Acanthaceae. Plantaginaceae.

7. Hubiinae : Fam. Rubiaceae^ ^fcaprifoliaceae. ValerianaceaeH
S.OCampanulinac Fam. Campanulaceae. Lobeliaceae.CT- Cucurbitaceae.^
:

9. Aggregatae Fam. Dipsacaceae. Compositae.

:

4
Class I. Gymnospermae ( )

Survey of the Development of the Sexual Generation

The flowers of the Gymnosperms are all unisexual. The macro-

sporophylls form the female, the microsporophylls the male flowers.
The two sexes are found either on the same individual (MONOECIOUS)
or each plant bears either male or female flowers (DICECIOUS). Leaves
forming an envelope around the group of sporophylls are found in
only a few flowers of the Gymnospermae.

The MALE FLOWERS

are shoots of limited length, the axis of which bears the

closely crowded and usually spirally arranged sporophylls. The scales which
invested the flower in the bud often persist at the base of the axis. The micro-
sporangia are borne on the lower surface of the sporophylls, two or more In- ing
present on each. Their opening is determined as in the sporangia of the Pteri-
dophyta by the peculiar construction of the outer layer of cells of the wall
(exothecium). The pollen - grains are spherical, and are frequently provided
with two sacs filled with air which increase their buoyancy and assist in
their distribution by the wind (Fig. 440 D).
A prothallium consisting of a few cells is formed on the germination of the
microspore. This lies within the large cell, which will later give rise to the pollen-
tube, closely applied to the cell wall. The prothallium is composed of from
1 to 3 cells. The first-formed cells (Fig. 441 A-C, v) correspond to the vegetative
cells of the prothallium. The spermatogenous cell (sp) which is cut oft' after these
divides later into the mother-cell of the antheridium and a sterile sister-cell (s)

adjoining the vegetative cells (Fig. 441 B, D). It is by the breaking down of this
latter cell that the antheridial mother-cell becomes free to pass into the pollen-
tube. On this taking place, or while the latter cell is still in its original position,
it divides into two daughter-cells (y) ; these are the generative cells or the male
sexual cells (Fig. 441 E). According to JUKL, Cupressus possesses a larger number of
 PHANEROGAMIA 481

D
FKI. 440. Pinus montawt. A, Longitudinal section of a ripe male flower (x 10). B, Longitudinal
section of a single stamen (x 20). C, Transverse section of a stamen (x 27). D, a ripe pollen-
grain of r: n an <ilre*tris(x 400).

. 441. -A', Pollen-grains

.-I
(iymnospcrms and stages in the development of the iK>llen-tuly
(if

A, Pollen-grain of Ginkgo enclosed in the sporangium (x 360).

Inlolxi still K, Pollen-grain of
l.ni-i
wropaea germinating on the tip of the niicellus ( x ItiO).
i ,

'/-A', Pinus ricio (after COULTER

and CHAMBERLAIN C, D x 300, K x 250), K, tip of a pollen-tulie which has reached the arche-
;

.-"iiiuni. f, Vi-.^etat ivi-. prothallial cell; gp. spermatogeiKius cell, which xi vl 's rise to the

;/, the two generative cells derived

aiithei-idial mother-cell (?n) and its sterile sister-cell (.<) ;
by
the division of the antheridial mother-cell ; nucleus of the pollen-tnhe, which in A" cannot be
/. ,

distinguished with certainty from the nucleus of the disorganised cells.

2 I
482 BOTANY

i. Ceratozamia longijolia.
442. Lonj:i- Fi<;. 443.--2VWM.V Ixtccat'a. Longitudinal section
tuilinal section
through a youni: ovule, through tli' sport igenous tissue, showing an
showing the nucellus (l), including a embryo-sac mother-cell which has undergi'iie
lar^e group of sporogenous cells (spor.) the t trail three of tlie fai'iiltat i\
division; >

and tin- integument (/) (After TREUB. niacri)s])oi-es aredegenerating, while the fourth
x 33.) is undergoing further development, (x 2">0.)

generative cells, while Taxus has

only one. In the Cycadinae and
Ginkgoinae the male cells appear as
spermatozoids, while in the other
Gymnospenns they are smooth cells
not provided with a cell wall.

In many Gymnosperms the

FEMALE FL4IWKKS or <.'<>NK*
resemble the male flowers in
being composed of an axis
bearing numerous, spirally ar-
ranged sporophylls. In other
cases they differ from this
type in various ways, which
will be described in the special

part below.
As a rule the macrosporophylls
bear two macrosporangia these;

consist of a nucellus and usually

a single integument. The nucellus
often becomes of large size by the
numerous periclinaldiviriona which
take place in the more superficial
layers of cells. In the same way the
sporogenous tissue, which origin-
ic. 444. Median longitudinal of an ovule
section ates immediately below the i-pider-
of I-icca culyans. e, Emliryo-.siic with the pro-
tilleil
n ,j Sj becomes deeply placed by the
thallium n, archegonimn showing ventral (o) and
;

'
rati<m of Iiumero u8 layers of
neck portion (c) ; >t, nucleus of e^'g-cell nr, nucellus; ;

p, i^iien-raius ; /, ix,iien-tui. :
t, lategomnti ,
cells -
Notwithstanding the con-
-Ainj.'. siderable size of the sporogenous
 SKIT. II PHANEROGAMIA 483

tissue only one, centrally placed macrospore-mother-cell is as a rule developed, all

the other cells remaining sterile. The mother-cell undergoes a tetrad-division (Fig.
443), and of the four resulting cells only one develops into an embryo-sac (macro-
spore). increases in size, first crushes its sister-cells and later the whole
This, as it

sporogenous complex of cells. Meanwhile, by the repeated division of the nucleus

and the macrospore becomes filled with the tissue of the prothallium
protoplasm,
(Fig. 444). The archegonia are formed at the apex of the prothallium ; each
consists of a large ovum and a short neck. As in the Pteridophytes a small ventral
canal-cell is cut off from the egg-cell shortly before fertilisation. Fertilisation takes

place by the entrance into the ovum of one of the generative cells, the development of

A B
,d

l-'ii.. 446. Picea ewcbd (.-I-', F). Finns larlc'w (D, K, fH-1). A, mature ovum with its nucleus (nn)
and the ventral-canal-cell (</); fi, the male nucleus (x) within the ovum : fusion of the
'
,

male and female nuclei D, the four nuclei produced by division of the nucleus of the embryo
;

have passed to the lower end of the ovum and are there in process of further division (only
two of the nuclei are visible in the section); K, four of the eight nuclei are contained in
independent cells, while the other four remain in the general cavity of the egg-cell F, further
;

division of the nuclei of the upper series (!, the cells of the lower tier have divided
; H, four ;

tiers of nuclei are present, those of the upper tier not being separated from the general cavity
of the egg-cell /, the elongation of the middle tier of four cells to form the suspensor (s) has
;

'

commenced. (A-<.' x 55, F x 110, after MIYAKE ; D, E, G-I x 104, after COULTER and
CHAMBERLAIN.)

which has already been followed this enters from the pollen-tube, which penetrates
;

between the cells of the archegonial neck. The nucleus of the fertilised ovum
results from the fusion of the male and female sexual nuclei (Fig. 445) and the

protoplasm is in part derived from the male cell. The development of the embryo
from the fertilised ovum presents great differences in the several orders and even
genera, and the following description applies to the species of Pinus.
By two successive divisions of the nucleus four nuclei are formed which pass to
tin- base of the
egg-cell, where they arrange themselves in one plane and undergo
a further division. Cell walls are formed between the eight nuclei of this eight-
celled pro-embryo. The cells form two tiers, those of the upper tier being in open
communication with the cavity of the ovum. The four upper cells then undergo
another division, and this is followed by a similar division of the four lower cells.
The PRO-EMBRYO thus consists of four tiers each containing four cells, the cells of
484 BOTANY PART n

the upper tier being continuous with the remaining portion of the ovum. In the
further development of the three lower tiers the middle tier elongates to form the
SUSPENSOR, pushing the terminal tier from which the embryo will arise into the
tissue of the prothallium or endosperm ;
the cells of the latter are filled with
nutritive reserve material.
In other genera a separation of the four rows of cells takes place, and each bears
a young embryo. As a rule, however, only a single embryo continues its develop-
ment in each macrospore although several archegonia may have been fertilised.
The embryo consists of a main root or RADICLE directed towards the micropylc.
an axis, the HYPOCOTYL, a whorl of from two to several COTYLEDONS and the
i-i.rMULE or bud of the future shoot.
The asexual generation which shows less uniformity will be described under the
separate orders.

Order 1. Cycadinae (
:>

This includes the single Family, Cyeadaeeae. These are woody

plants restricted to tropical and sub-tropical regions. The stem,
which undergoes secondary growth in thickness, is as a rule un-
branched and bears large, pinnate foliage leaves. These, which are of
firm leathery texture and persist for a number of years, alternate with
smaller scale-leaves and form a large terminal crown. The surface of
the cylindrical or tuberous stem is clothed with the scale-leaves and
the bases of the old foliage leaves. Mucilage ducts are present in
all parts of the plant. The vascular bundles are collateral but their
xylem consists of tracheides only.
The Cyeadaeeae are dioecious. Fig. 446 represents a female plant
of Cycas revoluta, in which the growing point forms alternate zones of
foliage leaves and macrosporophylls. When young the foliage leaves
are rolled up circinately as in the Ferns. One of the sporophylls is
represented in detail in Fig. 447. It shows the pinnate form of the
foliage leaf but is densely covered with hairs, and chlorophyll is
wanting. Towards the base two to eight macrosporangia are borne
on the margins. It is evident that each female plant of Cycas which
has reached the flowering condition exhibits a regular succession of
flowering and vegetative periods. The flower represented by the group
of sporophylls is always grown through by the further development
of the apex which does not branch. The male plant of Cycas and the
other Cyeadaeeae are morphologically more advanced in that they
regularly produce lateral buds. Their sporophylls are borne in terminal
cones often of great size, while the further growth of the plant is
effected by a lateral bud which continues the direction of growth of
the sympodial axis, displacing the cone to one side.
The cones consist of numerous oporophylls arranged spirally on
the axis. The microsporophylls bear large numbers of microsporangia
on the lower surface (Fig. 4 i8). The macrosporophylls of the cone-
bearing Cyeadaeeae are considerably modified as compared with Cycas,
 PHANEROGAMIA

and each bears two marginal macrosporangia (Fig. 449) ; only in

Stangeria are the macrosporangia situated on the lower surface.
The development of the sexual generation in the Cycadaceae differs in a number
of respects from the general scheme for the Gymnosperms as given above. The

Kit:. )4r>. fi/nix ri't-tilutit. ffinalf plant in Mmvrr. (Kroni ;i

iilioto^fapli.)

ditt'erences indicate that the group is more closely connected with tin-
Ptelidophyta,
and their nature will be evident from a description of Zamia floridann which \\u> ,

been fully investigated by Webber.

The female cones of Zamia bear numerous sporophylls, the hexagonal shirld-
sliaped terminal expansions of which fit closely together. Each sporophyll licars
a ji.-iir of
macrosporangia as is shown for (,V/Wn-.v////<V/ in Fig. 449. The macro-
2 I 1
 486 BOTANY

sporangium consists of the nucellus and an integument. The micropyle forms

iin open canal above the tip of the nucellus. At the period during which the male
cones are shedding their pollen, the macrosporophylls
become slightly separated from one another so that the
wind-borne pollen-grains can readily enter. A more or
lessextensive cavity (POLLEN-CHAMBER) has by this tiim
been formed at the apex of the nucellus, while the disin-
tegrated cells, together perhaps with fluid excreted from
the surrounding cells of the nucellus, form a sticky mass
which fills the mieropylar canal and forms a drop at it-

Kli;. 447. CarjM'1 of '

>cs re-

niltita. (After SACHS, re-

duced.) Fin. 44S. Stamen of 'i/cn* i-ii-i-iiiiilia, from U<-lo\v. (After RICHARD.)

entrance. The pollen-grains reach this drop and, with the gradual drying up ot
the fluid, are drawn through the mieropylar canal into the pollen - chamber.
During the development of the pollen-tube and the formation of the motile sper-
matozoids, which have been found in all the Cycads yet investigated, the embryo-
sac filled with the prothallial tissue is increasing
in size within the nucellus. As it crushes the
tissue of the upper portion of the nucellus it
approaches the base of the pollen-chamber. At
the apex of the embryo-sac are found the arche-
gonia, usually four in number, and separated
from one another by some layers of cells. Each
archegonium has a neck, and ultimately cuts
off a canal-cell. The archegonia are situated at
the base of a depression in the prothallium, the
archegonial chamber, which is. about 1 mm. in

depth and 2 mm. across. The jiollen-tubes grow

into this depression and liberate their sperma-
tozoids together with a drop of watery fluid in
which they swim. Since the bursting of the
is brought about by their contact
pollen-tubes
*
Fio. 440. CmUOKMria robiuto. Macro-
, , ,. ,. . .

s I(oro,,l,yll
wlth the projecting cells ot tin- neck, the sp-rn,,i-
with two macrosporania.
(After GOEBEL.) tozoids cannot easily lose their way to the egg-
cell. They require, however, to narrow consider-
ably in order to pass through the space between the neck cells. The nucleus
of the fertilised ovum soon divides, and the daughter-nuclei continue to divide

rapidly, until after the eighth division there are about 256 free nuclei within
the cell. These are crowded towards the lower end of the pro-embryo, where cell-
formation commences around them. The embryo is forced into the endosperm
SECT. II PHANEROGAMIA 487

FIG. 450. Formation of Spermatozoids in //(' fluri/lnnn. (i, Mature polleH7gratn (X 800);
v,vegetative prothallial cell the dark streak at its base indicates the position of another
crushed cell k, nucleus of the pollen-tube ; sp, spermatogenous cell. 6, c, rf, stages in the
;

development of the antheridium (h, c x 400 d x 200) v, persisting vegetative cell growing
; ;

into the sterile sister-cell of the antheridium (s) ; m, mother-cell of the antheridium, i.e. mother-
cell of the exine. In the mother-cell the large blepharoplasts (bl) which form.the
spermatozoids; e,

cilia are visible ;

in 6 and
they are star-shaped while in il they are composed of small granules,
c ,

which will form the cilinm-forming spiral band. Starch-grains are present in the pollen-tube,
and in c they are appearing in the vegetative cell and the sister-cell, both of which in d^are
packed with starch. In d the two spermatozoids (sj>) derived from the mother-cell are/seen
divided from one another by a wall. (After H. .1. WEBBER.)

Fn.. 4;">1. Upper end of the pollen-tube of /.// /u

floridana, showing the vegetative prothallial
cell (v), the sterile sister-cell (s), and the two
spermatozoids. o, Before movement of the
>l>errnatozoids has commenced 6, after
; the
beginning of ciliary motion ; the prothallial cell
is broken down and the separation of the two Fn.. 4.VJ. -xfum/'" jiiii'iiimin. Mature, free-

spermatozoids is taking place, (x circa 75. After swimming sperrnato/oid. (xlf>0. After
H. J. WEBBER.) H. .1. WEBBER.)

2 I 2
488 BOTANY I'AUT II

liy tin- elongation of a suspensor. It ulti-

mately' possesses a pair of large cotyledons, a

well -developed plumule between these and a
relatively short hypocotyl (Figs. 450-453).

Order 2. Ginkgoinae (
t;

The single representative of the Family of

the Ginkgoaceae which forms this order is
Hiiikijti This tree comes from Japan
tiiloba.

hut is often in Europe.

seen in cultivation
The long-stalked leaves resemhlc those of an
Adiantum, and are divided dichotomously into
two or more lobes. The llowers are dioecious.
The numerous stamens are situated on an
elongated axis which bears no enveloping
"
leaves. Microsporangia with an endotheciuin"
!']<;. -4.J3. /."mi" Jlnriilmiit. An ovum
(cf. Macrosporangia in pairs at the
p. 503).
immediately after the fusion of the
nucleus of a ipflmtnoU with the
summit of short shoots sporophylls reduced to
;

female nucleus has taken place. The a collar-like outgrowth around the base of the
ciliary kind of the s]>ermato/<>i<l
re-
sporangium (Figs. 454, 455).
mains in the upper portion of the pm- The development of the sexual generation
toplasm of the ovum. Aseconil spev-
inaU'/.oid has attempted to enter the
and the fertilisation is very similar to what has
OVUIP. (x 18, after H. J. WEBBER.) been described for Znmio. The fertilised ovum

Fi<:. 454.-';i;iJtyo biloba. Male branch with flower; the leaves an- iu.t yet full grown. 0, >>.

>. stone of same /, stone in cross section <7, in lonjd-

stamens ; c, female flower ./.
; fruit ;
: ;

tmlinsil section showin.u the emtiryo h. female (lower with an exceptionally lar-e Dumber
:

the
of ovules borne on separate stalks. (Male (lower and <-. nat. si/e ./, slightly
r.-<luce.) ;
;

i.thi-r ti-ui-s nia^iiiiied. AfVr RICHARD u-d after EICHLF.R.)

:
 PHANEROGAMIA 489

becomes filled with a continuous tissue and forms the embryo at the end farthest
from the micropyle. This grows out and
penetrates the endosperm, but there is no
clear distinction into suspensor and embryo.

7
Order 3. Coniferae ( )

The Coniferae include conspicuous

trees or shrubs with woody stems.
The possession of small, undivided,
firm leaves,flat or needle-shaped, and

usually lasting for several seasons,

is a common character of the
plants
of the order ; they are thus for the
Fio. 455. Longitudinal .section of a young
most part evergreen. All Conifers macroeporangium of (linkgo biloba, m,
are profusely branched, arid a distinc- micropyle ;integument p, pollen-
', ;

chamber e, embryo-sac w, outgrowth

; ;

tion into long and short shoots is of siwrophyll. The possession of a well-

usually evident. In many cases the developed pollen-chaml^r jxiints to the

direction and rapidity of growth of relationship with the Cycadaceac. (x

35. After COULTER and CHAMBERLAIN.)
the main axis differs from that of the
lateral branches. This is especially seen in young individuals ;
old
trees are often more irregular in outline.
The absence of vessels from the xylem of young plants and from
the secondary wood is an anatomical characteristic (cf. p. 134). Their
place taken by large tracheides with peculiar bordered pits on the
is

radial walls these form a very uniform wood.

;
The majority of the
Coniferae have resin abundantly present in all the parts of the plant.
The Coniferae in contrast to the Cycadinae are mostly inhabitants
of temperate regions, and are among the trees which approach nearest
to the polar regions. Within the tropics they are mostly confined to
mountains.
The Coniferae are divided into two families on account of differ-
ences in the floral structure.
The Taxaceae have female flowers with one or few macrosporangia ;
the latter are usually provided with an arillus. The flowers are
usually not definite cones. Mostly dioecious.
The Pinaceae on the other hand have a number of ovules in each
female flower, the latter being a cone with numerous sporophylls
borne on an axis. Arillus not present, usually monoecious.

Family Taxaceae. The plants belonging to this family are grouped in a

number genera distributed in the southern hemisphere.
of small The most
important genus is Podocarpus, the numerous species of which are widely distributed
in temperate East Asia and in Australia and New Zealand, and also occur as stately
trees on the mountains of the Asiatic tropics. The female flowers are small
shoots, the sporophylls of which are swollen and succulent one or two of them :

bear at the summit a single anatropous ovule surrounded by a fleshy arillus.

The male flowers, which are borne on the same or on distinct individuals, are small
490 BOTANY

cones consisting of numerous sporophylls attached to a short erect axis. Each

sporophyll bears two microsporangia on the lower surface the microspores are
;

provided with distended wings.

Taxus baccata is the only European representative of the family. The Yew,
which is now for the most part artificially introduced, had formerly a wide distribu-
tion as an evergreen undergrowth in our native woods (Figs. 456, 457). Isolated
examples of large size occur in North and Central Germany, and it occurs as a

Flo. 4a6. Taunts baccata.A, branch with feinalf dowers: '. two ovules mi the- s.-nnc shoot (nat.
with axillary, fertile shoot (x 2); f, median longitudinal section of a primaiy
size); B, leaf
and secondary shoot; r, vegetative cone of the primary shoot ; a,' rudiment of the aril; e,

nidiment of the embryo-sac n, nucellus i, integument; HI, micropyle (x 48). Fo/soxoi-s.

; ;

more important constituent of the vegetation in Switzerland, e.g. on the steep

slopes at Uetli. The Yew tree attains a height of 10 m. All the branches are
shoots of unlimited growth. The leaves stand on all sides of the ascending main
shoots but in two rows on the horizontally expanded lateral branches. They are
narrow, flat leaves and persist for several years. The tree is dioecious the flowers;

are situated on the lower surface of the twigs and arise in the axils of the leaves
of the preceding year. The male flowers are invested at the base by a number
of scale leaves and contain some 10 peltate stamens, each of which bears 5-9
pollen-sacs. The mode of opening of the sporangia is peculiar. The outer wall
 PHANEROGAMIA 491

splits at the base and along the side of each pollen-sac, so that the whole stamen
resembles an umbrella turned inside out the pollen remains for a time in the
;

pocket-like depressions, from which it is removed by the wind. The female

flower usually develops singly as a secondary, axillary shoot of the uppermost
scale leaf of a primary shoot the apex of the latter is displaced to the side and
;

does not develop further. Each flower consists of a single, atropous ovule with
one integument. The drop of fluid excreted from the micropyle of many Gymno-
sperms is especially well shown by the Yew. As the seed develops, a fleshy arillus

Km. 4.J7. Win* lii'n -lit". (.1 nut. si/f.) J'oiao.vofs.

.springs from base and surrounds the mature seed like a bright red cup.
its The
foliage and seed are poisonous, but the aril, which induces birds to distribute the
seed, is harmless.

Family Pinaceae. This family includes the most important

Coniferae, and on grounds of differences in leaf arrangement and in
the position of the ovules is divided into two sub-families. The
forms with the leaves opposite or in whorls are included in the
Cupressineae ;they also have the ovules erect. Cupressus, Thuja,
Juniperus, and Taxodium are representative genera ; the habit of
Taxodium is peculiar. All the forms with alternate leaves are included
in the Abietineae, and, almost without
exception, they also possess
492 BOTANY

inverted ovules. The genera Araucaria and Agatliis Sequoia and

Sdadopitys Abies, Picea, Larix, and Pinus, may be distributed in
three distinct groups which may be named Araucarieae, Sequoieae,
and Abietineae, in the narrow sense.

Sub-family Cupressineae. Some of the Cupressineae have needle-shaped

leaves in whorls (Juniper, Fig. 458), others have decussately arranged, scale-like
leaves (Thuja, Juniper UK siilinn. Fig. 459). 'The former type is to be regarded as

FIG. 458. Juniperus cum muni*. T\vij;.s bearing fruits and male flowers. (3 nat. size.) Of'f'li /.</..

tlie more primitive, for the seedlings ol Thuja have needle-shaped leaves, and
individual branches of scale-leaved forms of Juniperus revert to the needle-shaped
leaves in whorls of three. The short shoots of Taxodium have two ranks of leaves
and are shed as a whole.
The Cupressinae, with the exception of Juniperus, are monoecious. The male
flowers of Juniperus communis stand in the leaf axils. At their base are a number
of small scale-leaves (Fig. 460 a, a), above which come several whorls of peltate
sporophylls (c) bearing 2-4 microsporangia (d) on the lower surface. The sporangia
open by a vertical slit parallel to the long axis of the sporophyll. The female
flower.- occupy a corresponding
position. The scale leaves at the base (Fig. 460 b)
are succeeded by a whorl of carpels (c, b), each of which bears a single upright ovule
in a median position (c). After fertilisation a succulent parenchymatous growth
mainly of the basal portions of the sporophylls raises the seeds and presses them
together, without, however, obliterating the central spaee altogether. The three
 PHANEKOGAMIA 493

carpels become completely coherent above the seeds, but the place of union is still

Fie. 450. Juniperus Sabina : branch with fruit. 1'oisosoos.

indicated by the scar at the apex ot the ripe fruit the tip of each carpel is a
;

little back from the corresponding angle of the suture. The fruit takes two years

Fie;. 460. Juniperus com muni*, c, -Malt; flower; '-, fertile shoot witli female flower; c, female
flowerlwith one scale bent out of place; d, fruit. OFFICIAL. (After BERG and ScHMiDiyall
magnified.),

to ripen. The succulence of the carpels gives the fruit the appearance of a berry.
494 BOTANY

the only genus of the Cupressineae with such fruits ;

the others, such
as Cupressns, Thuja. Tit.riIit>i,,, have cones.
.luniperus cominnnis, Juniper, is a shrub or
small tree distributed over the northern hemi-
sphere. ,/. Xiil>in<t a
prostrate shrub of tin-
Alps and other mountains of central and southern
Kurope. The Cypress (f.'n/iri'sxus sempervircns)
in the Mediterranean region. 7'. /-///>/// //s-

>!<// n m is a deciduous tree, forming extended

swampy woods on the north coast of the Gulf of

Mexico from Florida to Galveston. T. mexicanum
is evergreen and is widely distributed on the

highlands of Mexico very large specimens occur ;

such as the giant tree of Tule, which at a height

of 40 in. was 30 m. in circumference, and was
fjl estimated by VON HrMuoi.ivr to be 4000 years old.

:<;. 401. flnamiUvettria. Ovulitcniu>

scale (//) bearing two ovules s and
the keel c. The bract scale /; is

visible l>ehind. The free margin of

the integument of the ovule forms
two prolongations (TO), (x 7.)

Sub-family Abietineae. The floral

structure of the Abietineae may In-
described in the first place. The male
flowers (cf. Fig. 440, p. 481) consist of
an axis bearing scale leaves at the base,
and, above this, numerous stamens :

the pollen -sacs (microsporangia) art-

situated on the lower surface of the
stamen. In the Abietineae in the
narrower sense there are two pollen-
sacs, but in Agathis and Araufurin
there are 5-15. The projecting tip of
the sporophyll varies in size and ap-
pearance, but is as a rule triangular.
The microspores are usually winged.
The female flowers are always cones.
consisting of an axis bearing the closely
approximated scales, which protect the
ovules the scales later become liqui-
;

fied. In Agathis and Araucaria each

scale bears a single
anatropous ovule at l-'n;. 4iVJ.--. -I /)(>.< )Tt;ii"tn. a. Male (lower; /, scale
its base.
(The cone-bearing Cupressineae leaves : 7i. .-.I'lu^pliyHs. /.. Cmii-. <, Carpel, viewed
have an outgrowth of the scale on from lielow (dor>al surface), hi>win.", the fertile

which the ovules are seated the out- and cover-scale rf, the same viewed from alwve
;
;

(ventral surface). (After BEU" and SCHMIDT:

growth not sharply marked off from
is
a, c, il, nat. si/i- : l>, n-duced.)
the scale.) The condition of affairs in
Sequoia and Sciadopitys is similar, but the outgrowth is more definitely defim-d ;

each scale bears 4-9 anatropous ovules. In the Abietineae proper the limits of the
 PHANEROGAM I A 495

two scales are still more marked. The two anatropwis ovules are borne on au inner
scale, which, at its Lase, is continuous with the scale of the cone. THE OITKI:
si .U.K. CAU.KH THE lUl.YCT SCALE, THE INNEK, THE OVULIFEKOUS SCALE (FigS.
IS

461, 462). The ovuliferous scale is the more strongly developed and it is the part
that becomes lignified and affords protection to the ovules. Even at the period
of flowering the bract scale is usually concealed by the ovuliferous scale and only
to be 'detected on close inspection. In other forms, however (e.g. Abies bracteata,

Via. 4ii3. 1'M'a aci-ilx* (i nat. si/e). 1, Twix with male tloneis. ', Terminal female flown-, j,
Pendulous cone. !,, MicTosiM)iij]>liyll. .", MaCKMporophyll ; tlie bract-scale is covered by the
larjje, bent-back, ovuliferous scale an ovule is visible at the base of the ovuliferous scale.
;

Kipe seed with the win;.: funned by a detacheil portion of the ovuliferous scale, (x 4-fi.)
'-',

A. /ii'ctiiinta, Fig. 462 b, /'*< in/u/sni/ti limujJasi-i, etc.), the bract scales even in the
older cone project prominently between the ovuliferous scales.
The view here followed, that the ovuliferous scale is a placental outgrowth of
the bract scale bearing the macrosporangia, and that its increase in size can be
traced in the series of the Araucarieae and Abietineae, is that supported by SACHS,
EICHLER, and GOEBEL. Attention must be drawn, however, to an alternative view
supported by STHASi;nif;Kii and CEUAKOV.SKY among others. According to this
the ovuliferous scale corresponds to the coherent bracteoles of an axillary shoot
borne in the axil of the bract scale. The cone itself would thus not correspond to
a single flower bearing a number of sporophylls, but to a system of shoots, i.r. it

would be an inflorescence.
496 BOTANY PAKT n

Most important Genera and Species. Ayathis'(Dammara) is distributed in

the Malayan Archipelago and extends to New A. australis and A.
Zealand ;

Dammara yield Kauri Copal but no Dammar Resin Araucaria brasiliana and A.
;

imbricata are stately, S. American, forest trees A. excelsa, A. Cookii, and other
;

species with very limited distribution are extensively cultivated as ornamental

trees. The genus Sequoia includes the most gigantic trees known specimens of
;

S. gignntca from the California!! Sierra Nevada attain a height of 100 m. and' a
diameter of 12 m.
European Forest Trees. Picea excelsa, the Fir (Fig. 463), is a fine tree of
pyramidal shape it has no short shoots, and the long shoots bear on all sides
;

pointed, quadrangular, needle-shaped leaves, which on horizontal or pendulous

branches stand more or less erect. Male flowers as a rule on shoots of the previous
year ;
on flowering they become twisted into an erect position. The two pollen-
sacs open by a longitudinal slit. Female flowers terminal on the shoots of the
previous year usually near the summit of the tree. They stand erect at the tiim-
of flowering. The ripe cones are pendulous and, after setting the seeds free from
between the scales, fall in pieces. The development of the seeds is completed in
one year. The male and female flowers occur on the same individual. Picen
orientalis from Asia Minor, and Picea alba from N. America are frequently
cultivated.
The Silver Fir (Abies pectinata, Fig. 462) is a native of the mountains of the
middle and south of Europe. It bears only long shoots. The flat, needle-like
leaves, marked below by two white lines and emarginate at the tip, are borne on all
sides of the axis, but are twisted- into a horizontal position on the branches
illuminated from above. The male flowers stand in the leaf-axils on the under
side or on the flanks of the shoot, and grow downwards so that the pollen-sacs are
directed upwards. The wall of the sporangium opens by an obliquely longitudinal
split, which gapes widely and allows the winged microspores to escape. The
female flowers arise from the upper side of a branch and are directed vertically
upwards. The bract-scales are longer than the broad, ovuliferous scales. The
fertilised cones retain the upright position, and when ripe the scales separate from
the axis and so set the seeds free from the plant. The development of the seeds
takes a year. Abies Nordmanniana from the Caucasus, A. pinsapo from Spain,
A. concolor, A. balsamea, and A. nobilis from N. America are in cultivation.
Larix europaea, tlie Larch, is one of the few deciduous Conifers and replace* its
foliage annually. There is a differentiation into long and short shoots. The
former bear the narrow linear leaves on all sides and continue the branching of the
pyramidal tree the lower branches of which often droop downwards. The short
shoots arise in the axils of the leaves of the long shoots of the preceding year,
and bear a rosette of 30-40 leaves which are somewhat shorter but resemble those
of the long shoots. The flowers occur in a position corresponding to that of the
short shoots. The male flowers are bent downwards when fully developed, and
the opening of the upwardly directed pollen-sacs occurs as in Abies. The erect
female cones produce seed in the same year.
The most advanced differentiation of the vegetative organs is found in the
genus Pinus P. silvestris, the Scotch Fir, will serve as an example (Fig. 464).
;

Young seedlings in the first or second year have long shoots bearing needle-shaped
leaves. On older plants this type of foliage is lost ; the needles are replaced by
colourless, membranous scale leaves in the axils of which stand the short shoots.
These have 2 in other species 3 or 5 needle-shaped foliage leaves. The latter
are about 5 cm. in length, their flat surfaces are turned to face one another and the
KM;. 464. J'i/ius silvestris(% n.it. size). 1, Shoot of unlimited growth hearing short slioots
; at the
top the shoot of the current year. At the oasejo the latter are numerous male flowers each in
the place of a short shoot, and nearer the tip. brown seal leaves, in the axil of each of which
is a short shoot. 2, Similar branch bearing a young female flower at the summit of 'the shoot
of the current year, in place of a branch of unlimited growth. Two dependent green cones
are borne on the shoot of.the preceding-vear. 3. Cone of thejyear before last, opened to allow
of the escape of the seeds. .',, A microsporophyll. f, Macrosporophyll from the adaxial side
showing the ovuliferous scale with the two ovules at the base. C, Macrosporophyll from the
abaxial side showing the small bract-scale below the large ovuliferous scale. 7, Ripe seed with
its wing derived from the superficial layers of the ovuliferous scale, (x 4-7.) OFFICIAL.
497 2K
498 BOTANY

outer or lower surface is convex. The growing points of tin- short shoots soon
become funetionless unless stimulated to activity by the death of the apical bud.
The male flowers appear in large numbers at the base of the long shoots of the
current year and stand in place of the short shoots. One or several female (lowers
arise at the tip of similar long shoots, and each corresponds in position to a shoot
of unlimited growth. At the time of flowering they are erect, but are bent down-
wards after pollination. The seeds ripen in the second year, and are set free by
the separation of the scales of the cone which till then have been closely pressed
together. The cones subsequently are shed (Fig. 464). Pinus montaim, a dwarf
Pine occurring on mountains P. pinea, P. ccmbra, with edible seeds P. laricio,
; ;

Corsican Pine from Austria; P. Pinaster, Maritime Pine from the Mediterranean
region ;
P. tacda, ]'. .SY/W,//*. Weymouth Pine, P. Lambertiana from N. America.
Cedrus. Cedars from the forests on Atlas and
Lebanon.
POISONOUS. J a Hi in riixSiiliiiui, Tii.i'aslmccata.
OFFICIAL. Pinus sylvestris and other species
produce OLEUM TEREBIXTHIXAE and RESIN A :

Abies balsamea supplies TEREBIXTHIXA CAXA-

Di.xsis P. palustris and P. taeda yield THUS
;

AMKRICAXUM ;
P. excelsa yields PIX BURGUXDICA;
P. sylvestris, etc. PIX LIQUIDA; P. piimilin. OLEUM
PIXI Juniperus oxycedrus and other species yield
;

OLEUM CADINUM Juniperus commit nis, OLEUM

;

JUNIPERI.
8
Order 4. Gnetinae ( )

The only Family in this order is that

of the Gnetaeeae, to which only three
genera belong ; Ephedra (Fig. 465) leaf-
less shrubs of warm dry regions of the
Northern hemisphere ; Welwitschia mira-
FK;. 465. Kphedm altissima. 1. Habit bilis, a monotypic plant from the deserts of
of a male inflorescence, 2. A South-West Africa the widely expanded
;
inflorescence with unripe ''

(5 nut. si/e.)
summit of the stem bears after the coty-
ledons only a single pair of leaves, which
are 1 m. in length and continue to grow at their bases Gm'tinu ;

(Fig. 466), tropical trees or climbers with pairs of broad, reticulately

veined leaves. These genera, while differing widely in appearance,
agree in possessing opposite leaves (in Ephedra reduced to scales),
in the development of vessels in the secondary wood, the absence
of resin canals, and in the presence of a perianth to the flowers which
are usually dioecious (Fig. 467).
These points of agreement with both Gymnosperms and Angio-
sperms make the group in many ways an intermediate one between
the two classes. The development of the sexual generation in the
three genera is as various as their external habit.

The microspores in their development and germination show no differences from

those of other Gymnosperms. The macrospores of Ephedra and Welwitschia have
 KII.. tiiti. Gnetum Gnemon. Hrancli with male inflorescences. (I, nat. size.)

--/

ptJL.

Fie;. I'll. ~.l. l-'.[ihi'ilra altlssima. Male flower, Fin. 4ti8. Gni'liiM Rumphiaunm. Apex
X 16; pg, perigone ; b, leaf. B, (!n>-tinn of the embryo-sac shortly before the
Gnemon, lon^ittiiliniil section of a female formation of the embryos, wk, female,
flower (x 3'2. After Lorsy) n, nuc-llu- ;
; nuclei ; mk, male nuclei ps, pollen- ;

ii. iiuifi. and ai, outer integuments y;i. : tube ; pk, disorganised nucleus of the
integument-like investment or perianth. pollen-tube, (x 250.)

499 2 K 1
500 BOTANY

well -developed protliallia ;

in the latter plant the archegonia are reduced to
elongated cells penetrating the tissue of the
nucellus. In G'netum, on the other hand, either
no prothallium is formed hut the embryo-sac
becomes tilled with protoplasm in which are
numerous nuclei, or a prothallium is developed
in the lower portion of the embryo-sac while
the upper portion contains protoplasm with
In t nuclei. Each of the two generative cells
from the pollen-tube fuses with a female nucleus.
Of all the fertilised cells resulting from the
penetration of a number of pollen-tubes to the
embryo-sac only one develops into an embryo
(Figs. 468, 469).

9
Fossil Gymnosperms ( )

In contrast to what was seen to be the .

asr
for the Pteridophyta, Gymnosperms have not
yet been detected in Silurian strata. They
appear first in the Devonian, but are sparingly
represented and first form an important con-
stituent of the flora in the Carboniferous. From
the Cycadofilices. which possessed stems with
secondary thickening and fern-like foliage and
V\t:. -ttiy.UiU'liuit
lineman. Embryo-sac. had been hitherto regarded as Pteridophyta,
r,upper portion with free nuclei y>. :
OLIVER and SCOTT (
9il
) have recently separated
lower portion filled with the tissue of
the Pteridospermeae ;
these may be briefly char-
the prothallium ps, pollen-tube
;
: .

two arrested embryo- sacs. (x (56,

acterised as fern-like spennaphyta. They have
after LOTSY.) si own that the seeds of Lagenostoma Lomaxi,
i

belong toLyyiuodtudron, oneof the Cycado-

\Vill,
filices with fern-like foliage, and that this plant must therefore be placed in the
Pteridospermeae. It is highly probable that the whole of the Lyginopterideae and
possibly of the Medulloseae also must be placed in this group. The structure of
the seed of Lagenostoma resembles that of Cycas, but, instead of the single thick in-
tegument of the latter, had a cupular investment to the seed on the outside of the
integument. In North America fragments of leaves referred to Cordaites have been
found. This genus belongs to a peculiar type confined to the Palaeozoic rocks. Owing
to the excellence of the preservation of their remains the morphology of the Cordaiteae
is as well known as that of the existing Gynuiospernis. They were lofty trees with
linear or broad and lobed leaves. Their flowers differ considerably from those of
recent Gymnosperms. An important fact as bearing on the phylogeny of the group
is the presence of a male prothallus as a small multicellular body. The ovules and
seeds show great structural agreement with those of the Cycadaceae.
With the exception of some less common fossils (Cycadites, Dicranophyllum},
the former of which is doubtfully referred to the Cycadinae, the latter more
probably to the Ginkgoineae, Cordaites is the most richly represented ty]ie of
Gymnosperm found in the Carboniferous rocks. Undoubted Cycadaceae make
their appearance in the lower Rothliegende. In later strata the genus Walchia, of
uncertain affinity, appears; the vegetative organs, which alone are known, recall
the lowest Coniferac (Araucariaceae). In the upper Rothlicgcndu undoubted
 PHANEROGAMIA 501

Ginkgoaceae (Baicra) make their appearance, with another type referred to the
Conii'erae
( Ullmannia).

The Cordaiteae disappear in the Rothliegende. The Gymuosperm flora can be

followed througli the Trias, in which it consisted of extinct types of Cycadineae,
Ginkgoineae, and Coniferae (Voltzia, in the Bunter Sandstone), to the Jurassic
period. In the latter it attained a great development in that both the Ginkgoineae
and the Cycadineae attained their maximum development and were represented
by numerous forms. True Araucarieae also appear in the Jurassic ;
on this
account, as well as on account of their organisation, this group may be regarded as
the oldest of the existing Coniferae.
In the Wealden Cycadineae and Ginkgoineae, along with some Coniferae, were
dominant among the Gymnosperms. On passing to the Cretaceous strata the
ancient types are found to be reduced, while the Coniferae become more numerous.
Among the latter existing genera appear (Dammara, Sequoia, Pinus, Cedrus,
Abies, Callitris, etc.). The Taxaceae also appear to be represented, but the remains
are of uncertain affinity.
The Tertiary Gymnosperms belong entirely to existing types and for the most
part to existing species. The Coniferae are dominant the Ginkgoineae are repre-
;

sented only by Ginkgo biloba, but this occurred in Europe along with other species
now limited to Eastern Asia or North America (Cryptomeria japonica, Taxodium
distichum, T. hetcrophyllum, Sequoia yiyantca, S. sempervirens, Pinus Strobus
etc.). One Cycadaceous plant (Enccphalartos) is also known.

CLASS II

10
Angiospermae ( )

The differences between gymnospermic and angiospermic plants

are well and shortly expressed by saying that the seeds of the former
are naked, while those of the latter are enclosed in a cavity. Only
a detailed account of the structure of the angiospermic flower will,
however, make the relations existing between the two great classes
of Phanerogams clear.
The Angiospermie Flower. The male and female .sporophylls of
Gymnosperms were borne in distinct flowers or even on different
individual plants, and the flowers had no special envelope. In
Angiosperms, on the other hand, a union of the two sexes in the one
flower which is thus HERMAPHRODITE and the investment of the flower
by coloured leaves (differing in appearance from the foliage leaves)
forming a PERIANTH is the rule. In contrast to the UNISEXUAL or
DICLINOUS flower with the sporophylls arranged spirally on an
elongated axis, which is characteristic of the Gymnosperms, the
perianth leaves and sporophylls in the Angiosperms are usually borne
in whorls on a greatly shortened axis. THE ARRANGEMENT OF THE
FLORAL LEAVES IN WHORLS, THE COLOURED PERIANTH, AND THE
II KRM APHRODITE NATURE OF THE FLOWERS ARE THUS CHARACTERISTIC
502 HOT ANN

OF ANGIOSPERMS, although these features do not apply without

exceptions to all angiospermic flowers. These differences depend
on the important factor of the MEANS OF POLLINATION. When,
as is the case with the Gymnosperms and the catkinate flowers
of Angiosperms, this func-
NN
tion is
performed by the
^ wind, the elongation of the
axis and the absence of an
investment of leaves around

Kn.. 471. Flourr ol A'-urus Calamus,

Fio. 470. Flower of J'ueonia peregrina, in longitudinal pg, pprigont- : c, androecinn; g,:

section, k and c, Perianth a, androecium g, gyn-

; ; ji.vnaeceum. (Enlanj"*!. Att'-r

aecetim. (J nat. si/t>.) ENOLER.)

the female receptive organ are advantageous. When, on the other hand,
pollination is effected by insects or birds, the conspicuousness given
by the presence of a perianth and other attractions, such as scent or
sweet-tasting substances, are necessary. The form of the flower, the
arrangement of the sporophylls in it, and the
place at which nectar is secreted must be
adapted to the
visiting insects (cf. p. 308).
It is to this, that the variety of form arid colour
exhibited in the flowers of Angiosperms must be
ascribed. According to the means of pollina-
f- tion, anemophilous, entomophilous, ornithophil-
ous etc. flowers are distinguished.
The perianth (Figs. 470, 471) consists of
two whorls of members these may be similar
:

in form and colour, when the name PERIGOM:

is given
& to them, or may be differentiated into
.4 and B, anterior
FII;. 472. j u r i
an outer green CALYX and an inner whorl or
and postal ior view of a sta-
men of Hyostyaihiis niger coloured leaves, the COROLLA.
; In every COD1-
/, the filament p, anther
p] ete fl ow er two whorls of stamens or micro-
; ;

c, connective (magnili.-d). , , ,

sporophylls come next within the perianth, and

within these again a whorl of carpels or macrosporphylls. The whorls
alternate regularly with one another. The stamens collectively form
the ANDROECIUM, the carpels the GYNAECEUM.
Each stamen consists of a cylindrical stalk or FILAMENT and of
the ANTHER the latter is formed of two THEC^E or pairs of pollen-
;
 PHANEROGAMIA 503

sacs joined by the continuation of the filament, the CONNECTIVE

(Fig. 472). According to whether the thecse are turned inwards,
i.e.towards the whorl of carpels, or outwards, the anther is described
as INTRORSE or EXTRORSE. The opening of the ripe theca depends
as a rule (except in the Ericaceae) on the peculiar construction of
the hypodermal layer of the wall of the pollen-sac. This is called the
fibrous layer or ENDOTHECIUM. On the other hand, in the Gymno-
sperms (excluding Ginkgo, 488), as in the Ferns, the dehiscence
cf. p.

is effected by means of the external layer of cells (exothecium).

As a rule the septum between the two pollen-sacs breaks down, so
that they are both opened by the one split in the wall. The
microspores in anemophilous plants are smooth, dry and light, and
adapted for distribution by the wind. In entomophilous flowers,
on the other hand, the exine is frequently sticky or provided with

KICJ. 474. Pollen-grain of Ml < xil cestris.

N, Spinous projections of the exine s, ;

Km. 473. .1, I'ollt'ii-^niin of L'ucurliita 1'epo (x 240) ; vertically striated layer of the exine ;

D, section of polleM-^niiii of Citcurbita vefruni.--n. p, the same seen from above n, places;

Allowing one of the lid-likf ar^as through which of exit of ]>ollfii-tul>fs. (After A.
the pollen-tubes protrude (x 540). MEVER.)

spiny projections, and the pollen-grains are thus enabled to attach

themselves better to the bodies of the insect visitors. They also
differ from the pollen-grains of the Gymnosperms in having more
or less numerous spots in the wall prepared beforehand for the
emission of the pollen-tube (Figs. 473, 474). Various modifications
of the androecium result from the cohesion, and branching of the
stamens, and will be described in the special part. Sterile stamens
which do not produce fertile pollen are termed STAMINODES.
The flower is terminated above by the GYNAECEUM. The CARPELS
composing this may remain free and each give rise to a separate
fruit (APOCARPOUS GYNAECEUM) or they unite together to form the

ovary (SYNCARPOUS GYNAECEUM). The carpels, as in the Cycadaceae,

as a rule bear the ovules on their margins, on more or less evident

outgrowths which are termed PLACENTAS. In apocarpous gynaecea

the ovules are thus borne on the united margins of the carpels,
each margin bearing a row of ovules (Fig. 476). This is termed the
f>04 BOTANY

VENTRAL SUTURE, while the midrib of the carpel forms the DORSAL
SUTURE. In syncarpous ovaries the ovules are similarly borne on the
margins of the coherent carpels. The placentation is termed PARIETAL
when the placentas form projections from the inner surface of the
wall of the ovary (Fig. 477 D). If the margins of the carpels project

Fm. 475. jtlthtifa rtjlii-iiuili*, Hower cut through KH.. 47(1. Transverse section of an ovary of
longitudinally ; a, epicalyx : l>, calyx ; c, Delphinium Ajatis, showing ovule placed
corolla ; (?, androecium. (After 13ER<; and horizontally; .--, ovule; p, placenta; o,
SCHMIDT, magnified.) ; wall of ovary ; r. vascular bundles, (x 18.)

farther into the ovary, and divide its cavity into chambers or loculi,
the placentas are correspondingly altered in position, and the
placentation becomes AXILE (Fig. 477 B). In contrast to such TRUE
SEPTA, formed of the marginal portions of the carpels, those that
arise as outgrowths of the surface or sutures of the carpels, as in
the Cruciferae, are called FALSE SEPTA. By the upgrowth of the

Flc. 477. Transverse sections of ovaries. A, l.olirlia ', /.'./'/"; >/; A7c/n</, /'
' '.

D, 1'as.fiflinii ; t il. placenta ; , ovules. (After LE MAOUT and DKCAISXK.)

floral axis in the centre of the ovary what is known as FREE CENTRAL
PLACENTATION comes about. The projecting axis cannot be sharply
distinguished from the tissue of the carpels. The septa, which
were originally present, are arrested at an early stage of develop-
ment or completely disappear, so that the ovules are borne on the
central axis covered with carpellary tissue and enclosed in a wall
formed by the outer portions of the carpels.
 PHANEROGAMIA 5or>

Each carpel usually is prolonged above into a a stalk-like STYLK

Via. 478. Different forms of gynaecea. A, Of Aconitum Napellus ; B, of Linum usitatissimum ; C, of

Nicotiana rustica ; D, style and stigma of Achillea Milltfolnnn f, ovary g, style H, stigma.
; ; ;

(After BERT; and SCHMIDT, magnified.)

terminating in a variously shaped STIGMA. The stigma serves as the

FIG. 479. Ovaries containing 1m sal ovules shown FKI. 480. Ovary of t'orniculum officlnale
in longitudinal section.A, Fagopyrum esctib 'utum with pendulous ovules, in longitudinal
(atropous) ; B, Armeria maritima (anatroi>ons). section. (After KERC, and SCHMIDT,
(After DUCHARTRE. x 20.) magnified.)

receptive apparatus for the pollen, and in relation to this is often

FK;. 481. Diagram of (.-I) liyixi.nynous (/-', L"), iTigynous, and ('') epigynous flowers.

papillate or moist and sticky (Fig. 478 D). When the gynaeceum
is completely syncarpous, it has only one style and stigma. In
506 BOTANY

Fig. 478 an apocarpous (A) and a syncarpous gynaeceum (O) are

represented, together with one in which the carpels are coherent
below to form the ovary while the styles are free (B).
The POSITION OF THE OVULES WITHIN THE OVARY may be erect,
pendulous, horizontal, or oblique to the longer axis (Figs. 479, 480).
In anatropous ovules the raphe is said to be ventral when it is

Fi<;. 4S2. Hypogyirnis HUWIT nf Humim-ulits .vvfccii/im with iiuim-riius, Mi|>crior ovaries home
upon a club-shaped receptacle. (After HAII.LON, magnified.)

turned towards the ventral side of the carpel, and dorsal if towards
the dorsal side of the carpel.
The differences in the form of the floral axis, which involve changes
in the position of the gynaeceum, lead to differences in the form of
the flower itself. Some of the commonest cases are diagrammatically
represented in Fig. 481 A-C. The summit of the floral axis is

I'M:. 4>3. Different flowers belonging to the family Rosaceae, cut through longitudinally. 1,
I'otentilla palustris,hyiwgynous 2, Alchemilla ulpim'. )>erigynous ; 3, Pyrus Mnlus, 'epigynous.
;

(After FOCKE in Nat. Pflanzen-familien, magnified.)

usually thicker than the stalk-like portion below it is often widened ;

out and projecting, or it may be depressed and form a cavity. If the

whorls of members of the flower are situated above one another on a
simple, conical axis THE GYNAECEUM FORMS THE UPPERMOST WHORL
AND IS SPOKEN OF AS SUPERIOR, WHILE THE FLOWER IS TERMED
HYPOGYNOUS (Figs. 482, 483). If, however, the end
of the axis is

expanded into a flat or cup-shaped receptacle, an interval thus separat-

ing the androecium and gynaeceum, the flower is termed PERIGYNOUS
 PHANEROGAMIA 507

(Fig. 483, 2). When the concave floral axis, the margin of which
bears the androecium, becomes adherent to the gynaeceum, the latter
is said to be INFERIOR while the flower is described as EPIGYNOUS.

The regions of the axis or of other parts of the flower which

excrete a sugary solution to attract the pollinating animal visitors
are called NECTARIES. Their biological importance is considerable.
In a typical angiospermic flower the organs are thus arranged in
five alternating whorls, of which two comprise the perianth, two the
androecium, while the gynaeceum consists of one whorl. The flower
is PENTACYCLIC. The number of members is either the same in each
whorl (e.g. three in a typical Monocotyledon flower, or five in a typical
Dicotyledon flower), or an increase or decrease in the number takes
place. This is especially the case with the whorls composing the
androecium and gynaeceum.
Some of the more frequent deviations from the floral structure described above

may be referred to here. Variation in the number of whorls is common. A

diminution in the number may result from the absence of one kind of sexal organ
in diclinous flowers (e.g. Myris^ica}, or from the absence of one of the two wh'orls

B
Fio. 484. A, ActinoniorpHic flower of Ueranium sangniin-uni. U. Zygoinoi pliic flower of
Viola tricolor. '
Asymmetrical flower of Canna indica.
,

composing the androecium or perianth (Compositae). Flowers in which the

androecium consists of only one whorl are termed HAPLOSTEMONOU.S, those with
two whorls of stamens DIPLOSTKMONOUS. On the other hand, the number of whorls
may be increased, and this may affect the androecium (A'osa), the gynaeceum (Punica
gra-natum), or the perianth (Hayenia abyssinica). Another simple modification
concerns the position of the whorls. The outer whorl of the androecium, in Erica,
for instance, does not alternate with the segments of the corolla but the stamens
stand directly above the latter. Such an androecium is distinguished as OBDIPLO-
STEMONOUS from the ordinary DIPLOSTEMONOUS type.
Variations in floral symmetry are more important both biologically and with
regard to the general habit of the flower. Those flowers which can be divided
into similar halves by more than two planes passing through the axis are termed
RADIALLY SYMMETRICAL or ACTiNOMORPHic (Fig. 484 A}. When a symmetrical
508 BOTANY PART n

li vision r,in only be made by two such planes of section the flower is BISYMMETKICAL
or BILATERAL belonging to the Fumariaceae). From the originally
(c.<j. />t\'iitt,'(i,

radial type the PORSIVEXTUAL (SYMMETRICAL, ZY<;OMORPHIC), and the A^YM-

MF.Tiii' A type> .-an 1- derived by the unequal si/e of ^nme of tlie inembers or
i.

tlieir displacement or suppression (Fig. 484 H, C cf. p. 16). In zygomorphic ;

flowers there is only one plane of symmetry by which the flower can be divided
into corresponding halves while in asymmetrical flowers no such plane of symmetry
;

exists. Dorsiventral flowers tend to take up a particular position in relation to

the directive force of gravity and exhibit a high degree of adaptation to insect
pollination. According to whether the plane of symmetry coincides with the
MEDIAN PLANE of the
flower (i.e.plane the
passing through the axis
of the flower and the main
axis), forms an acute
angle with the median
plane, or is at right
angles to it, flowers are

distinguished as MED-
IANLY DORSIVEXTRAL,
OBLIQUELY DORSIVEX-
TRAL, Or TRANSVERSELY
IMIRSIVEXTRAL res)
-

tively (Fig. 485). Radi-

Fl<!. 485. Flowers of, A, SiiitiUm-in nl/iiiia, iiiiMlianly xyj, n-

r

morphic(ilorsivi-ntral) ; /;, .If'gtxilvjhiiipocattainivi, obliquely

xynoinorpliie (tlorsiventral) ; '', I'oryilalis Intea, tiiinsvi-isi-ly Fn.. -!'>. Diagram of a penta-
y.yX'iinorpli if (dorsi ventral) ; il, hrart-. cyclic flower (Lilium).

ally symmetrical monstrosities of normally zygomorphic flowers are termed

I'ELORIC.
If a diagram of the arrangement of the members as they are seen in a crn-s
section of an opened flower-bud is constructed (p. 16) and so oriented that the
transverse section of the axis of the inflorescence stands above, that of the bract
In-low the ground plan of the flower, what is known as a FLORAL DIAGRAM is obtained.
The accompanying floral diagram (Fig. 486) is that of a monocotyledonoiis flower
with live whorls of members. A FLORAL FORMULA gives a short expression for the
members of a flower as shown in the floral diagram. Denoting the calyx by K,
the corolla by C (if the perianth forms a. perigone it is denoted by P), the androe-
cium by A, and the gynaeceum by G, the number of members in each case is
placed after the letter. Thus the floral formula of the monocotyledonoiis flower
represented in Fig. 486 would be, P3 + 3, A 3 + 3, G 3. When tin-re is a large
 PHAXEROGAMIA 509

number of members in a whorl the symbol oo is used denoting that the number is

large or indefinite. Such a formula may be further made to denote the cohesion
of the members of a whorl by enclosing the proper number within brackets, and by

placing a horizontal line below or above the number of the carpels the superior or
inferior position of theovary is expressed. By placing an arrow before the formula
the dorsiventrality of the flower may be indicated, and by varying the direction of
the arrow it can be made clear whether the dorsiventrality is median, oblique, or
transverse. The formula for the Lily given above and for a number of other
flowers would thus take the following forms :

P3 + 3, A 3 + 3, G (3).
Lily.

Buttercup. K 5, C 5, A oc G oo ,
.

Laburnum, j K5, C5, A (5 -1-5), Gl .

Hemlock. K 5, 05, A 5, G(~2).

Artemisia. K 0, C (5), A (5), G (2).

Inflorescence. In many Angiosperms the single flowers are borne in large

numbers on special branch systems which differ in anumber of respects from the
vegetative branch systems and are
termed INFLORESCENCES. In inflores-
cences, as in the vegetative region (cf.
p. 20), branches usually arise only
from the leaf-axils, the subtending
leaf being termed a BHACT. Leaves
borne on the axis of the flower itself
are known as BRACTEOLES. Bracts
and bracteoles may be classed to-
gether as BRACTEAL LEAVES (Fig.
487). The MEDIAN PLANE passes
through the axis and the middle line
of the bract. The bracteoles, and
the branches produced from their Kll
.

4S7 ,.-,,,,., .,,, ,

,.,,/ .,/; with bract
axillary buds, may lie in this median (</) ami biacteoles (*).

planewhen they are said to have a

MEDIAN POSITION, or more commonly they stand right and left of it, and their
position is described as TI: VNSVKKSK (Fig. 487).
According to the relative development of the main and lateral axes in an
is made between RACEMOSE inflorescences, in which the
inflorescence a distinction
main axes are strongly developed and exceed their lateral shoots, and CYMOSE
inflorescences in which the lateral axes are more strongly developed and overtop
the main axis. The former may be also termed MONOPODIAL in contrast to
the SYMPODIAL or cymose type. The more frequent kinds of inflorescence are
enumerated below.

I. Racemose Inflorescences.

(rt) Lateral axes unbninched.

1. RACEME stalked flowers borne on an elongated main axis (Figs. 488 A, 489).
;

2. SPIKE; flowers sessile on an elongated main axis (Figs. 488 B, 490). A

spike in which the axis is thickened and succulent is termed a SPADIX a ;

spike which, after flowering or after the fruits have ripened, falls off as a
whole, is a CATKIN (Fig. 491).
3. UMBEL flowers stalked, on a shortened main axis (Figs. 488 C, 492).
;
510 BOTANY PAKT II

4. CAPITULUM or HEAD ;
flowers sessile, on a shortened main axis (Fig. 488 D).
(b) lateral axes branched.
f>. PANICLE ;
a main axis bearing racemes laterally (Figs. 488 ,
l'.
|; ! .

Flo. 488. Diagrams of racemose infloresceiio's. .1. Uitri-mi'. /.'.

S|>ikf. '. Uinl'l.
D, Capitnliim. A', Panicle. F, Comiwund umbel.

6. COMPOUND UMBEL ;
an umbel bearing small umbels in place of the single
flowers (Fig. 488 F).

II. Cymose Inflorescences.

(a) The relatively main axis bearing more- than two lateral branches ;

PLEIOCHASIUM.
 PHANEROGAMIA 511

Fn;. 491. Catkin of Corglus

americana. (After DD
CHARTRE.)

Kic. 4!Ml. Spike of l'/i',it<iii<i

laneeolutn. (After I*r-

CHARTRE.)

Fl<i. 4S9. Racenii- of l.hturin

striata. i{, bracts.

Fi<;. 4!i Umlx'l of the Clii-rry.

(After DUCHARTRE.)

Fn;. 4!>3. Panicle of Y turn filament

[(Reduced.)
512 BOTANY

(b) Each relatively main axis bears two lateral branches ; DICHASIUM (Figs. 494 K,
495).
(c) Each relatively main axis bears one lateral branch MONOCHASITM.
;

1. The successive lateral axes all fall in the median plane.

(a) all the branches arise on the same side DREPAKITM (Fig. 494
; C, D).

7"
J(

6(
*
 PHANEROGAMIA 513

The Development of the Sexual Generation in the Angiosperms ("). On

germination the MICUOSPORES of Angiosperms form an antheridial mother-cell
(Fig. 497 m) which is clearly de-
limited from the large pollen-
tube cell, but is not enclosed
by a cell wall. The antheridial
cell passes into the pollen-tube,
and sooner or later divides into
two generative cells (Fig. 497 g).
The nucleus of the pollen-tube
(k) is usually visible in the
neighbourhood of the gener-
ative cells. The absence of the
small prothallial cells, and of
a sterile sister-cell of the au-
theridium, as well as the
absence of a cell wall from the
mother-cell of the autheridium,
are points in which the Angio-

sperms differ from Gymno-

sperms. The reduction of the
male prothallium has thus gone
so far that only the indispens-

ably necessary parts remain.

The characteristic differ-
ences which the Angiosperms
show from the general course
of development of the MACRO-
FIG. 495. Cyrnose inflorescence (dichasium) of Cerastium
SPORANGIUM in the Phan-
colllniiin. t-t"", successive axes. (After DCCHARTRE.) erogams commence with the

FIG. 49". Pollen-grain of Lilium Martagon and its

germination, k, Nucleus of the large vegetative
cell of the pollen-tube m, small mother-cell of
;

Fit;. Ut>. Scorpioid cyme of

Symphytum the antheridium ; g, generative rolls. (After
aspfrrimum. (After DUCHARTRE.) GUIONAHD. x 375.)
2 L
SI 4 BOTANY

cell divisions in the single, functional macrospore- mother -cell resulting from
the tetrad division (Fig. 498). The "n:iM.\r.Y NUCLKUS
of the embryo-sac"
divides and the daughter-nuclei separate from one another. They divide twice
in succession so that eight nuclei are present. After this cell -formation commences
around these nuclei. Both at the upper or micropylar end of the embryo-sue
and at the lower end three naked cells are thus formed. The two remaining
"ror.Ai: NUCLEI" move towards one another in the middle of the embryo-sac,
and fuse to form the "SECONDARY NIVLKUS of the embryo-sac." The three cells
at the lower end are called the ANTIPODAL CKLLS they correspond to tin :

Fio. 498. Development of tin- embryo-sac in l'nlii<j<iii<tin liirnfinitnui. M. Mother-cell of the

embryo-sac: <>ni>, embryo-sap; \ft, sterile sister-cells; c, cg^-rell ; .. syncrnid:c :
j>, jwlar-
nnclei ; a, antipodal cells; k, secondary nucleus nt tl inbryo-sac i-hn. clialaxa ; : ////'. micro-
pyle ; ni, ii. outer and inner integuments /I/H. funicultis.
; (1-7 x 320, 8 x I'.i'i.)

tive prothallial cells, which in Gymnosperms fill the 'cavity of the embryo-sac,
and in Gnetum Gneinon form a mass of tissue filling the lower end of the
latter. The three cells at the micropylar end constitute the EGG- APPARATUS.
Two of them are similar and are termed the SYNERGID^E, while the third, which
projects farther into the cavity, is the EOG-CELI, or OVVM itself. The synergidse
assist in the passage of the contents of the pollen-tube into the embryo-sac. Here
also the process of reduction has gone as far as possible ; in place of the more or
less numerous archegonia of the gymnospermous macrospore only a single egg-cell
is present. The significance of the synergidse is difficult to determine unless tliey
are regarded as archegonia which have become sterile.
The microspores, which cannot reach the macros]>ore directly, germinate on tin-
stigma. The pollen-tube penetrates for the length of the style and as a rule the
 SECT. II PHANEROGAMIA 515

tip enters the mieropyle of an ovule and so reaches the apex of the nucellus. Of
late years a number of exceptions to this usual course have become known. Treub
first showed in Casuarina that the pollen-tube entered the ovule by way of the
chalaza, and thus reached the peculiar sporogenous tissue, which in this case
develops a number of embryo-sacs. CHALAZOGAMY, as this mode of fertilisation is
termed in contrast to POUOGAMY, has no special systematic significance, since it is
found to occur in numerous and unrelated plants. As NAWASCHIN has shown in

Kit;. -I'.'li. Ovaiv ( Polygonvm I'^

vulus (luring t'tTtilisation. fa, Stalk-

like base ol n\:iry /, fnilictllu.s ; chit, :

chalaxa ;
*u, niicellns mi, mirropyle
; ;

ii. iniiei
1

.
i' . outer integument; e, Fifi. 500. Ovule of Ulmun i'ihinculatti. <, Kmbryo-
embryo-sac ; ek, nucleus of embryo- sac ; in, iiiici-n|iylf ; //, diala/a : t, |Hicket-like .s))aci'
KOC ; c), ege;-appiiratus ; mi, antipodal between the inte^iiiiients. Tin 1
jKdlen-tulx', ]ix, pene-
('11s; ;;. style; n. stijjma; />. l>ollen- trates dii-fctly tliiDux'U the two integuments and
Krains ; /, pollen-tubes, (x 48.) reaches the apex of the nucellus. After NAW \scnis.) (

the case of Ulmus, the pollen-tube in all these plants attempts to avoid growing
across hollow cavities, possibly because it needs to be continually nourished at the

expense of the surrounding tissue. A fuller understanding of chalazogamy and

related phenomena is approached if they are regarded from this biological point of
view (Figs. 499-501).
When the pollen-tube, containing the t\vo geiirmtivr o-lls. h;is readied the
embryo-sac, its contents escape and pass by way of one of the synergidae to the
ovum the corresponding synergida then dies. One of the two generative nuclei
;

fuses \\ith the nucleus of the ovum, which then becomes surrounded by a cellulose
wall. The second generative nucleus passes the ovum and unites with the large
516 BOTANY

secondary nucleus of the embryo-sac to form tin- EXDOSPEKM NUCLEUS (Figs.

502-504). Both the male nuclei an- often spirally curved like cork-screw, and
;i

N AW ASCII ix, who tirst demonstrated the behaviour of the second generative nucleus,
compares them to the sperinatozoids of the Pteridophyta.
The further develop-
ment usually commences by the division of the endosperm nucleus, from which a
the wall of the embryo-sac
large number of nuclei lying in the protoplasm lining
are derived. The endosperm arises by the formation of cell walls around these
nuclei and their proper surrounding protoplasm, and by the increase in number of
the cells thus formed (Fig. 95) to produce a massive tissue. The distinctive feature
of the development of the endosperm in Angiosperms from the prothallus of

Flo. 502. Funkia nrntn. Apt-x of nucellus,

Fio. 501. Longitudinal section of -an ovary of showing part of embryo-sac and egg-ap-
Juglans regia to show the chalazogamy. ps, paratus. A, Before, B, during fertilisation ;

jiollen-tube e, embryo-sac
; ; cha, chalaza. o, egg-cell; s, synergidii' t, pollen tube
; ;

(Somewhat diagrammatic, x 6.) n, nucellus. (x 390.)

Gymnosperms lies in the interruption which occurs in the process in the case of

the endosperm. In the embryo-sac, when ready for fertilisation, only an indication
of the prothallus exists in the vegetative, antipodal cells. The true formation of
the endosperm is dependent on the further development of the embryo-sac, and
waste of material is thus guarded against. The starting-point of this endosperm
formation is given by the secondary nucleus of the embryo-sac, which needs to be
stimulated by fusion with the second generative nucleus (the so-called vegetative
fertilisation) to form the endosperm nucleus, before it enters on active division.
From the fertilised ovum enclosed within its cell wall a PKO-EMBRYO consisting
of a row of cells is first developed the end cell of this row gives rise to the greater
;

part of the EMBRYO. The rest of the pro-embryo forms the SVSPEXSOU. Between
the embryo and suspensor is a cell known as the HYPOPHYSIS, which takes a small
 PHANEROGAMIA 517

part in the formation of the lower end of the embryo. The segmentation of
the embryo presents differences according to whether the plant belongs to the
Monocotyledons or Dicotyledons. IN THE LATTI.U TWO COTYLEDONS ARE FORMED
AT THE END OF THE GROWING EMBRYO (FlG. 505), AND THE GROWING POINT OF
THE SHOOT ORIGINATES AT THE BASE OF THE DEPRESSION BETWEEN THEM.
MONOCOTYLEDONS, ON THE OTHER HAND, HAVE A SINGLE LARGE TERMINAL
COTYLEDON, THE GROWING POINT BEING SITUATED LATERALLY (Fig. 506).
In both cases the root is formed from the end of the embryo which is directed
towards the micropyle ;
its limits can bo readily traced in older embryos, and it

I .
508. Ulinni Martagon.
Fertilisation <>{ Fl<;. .">04. -.1, Kmlii vo-sar of lli-lin ntli .< mi n u >.'x

One of the male nuclei

is close to the (alter NAWASCHIN). H. The male nuclei more
nucleus of tin' ovum, the other is in con- highly magnified. ;>x, iiolli-n-tulie ;
,vj, &,,

tact with the nuclei of the embryo-sac. vvnerjji'lse ; </>[, .-'/'>. male nuclei ;

(After GncXARD.) cell ; ek. nucleus of embryo-sac ; .

antipodal
cells.

can be seen that a few cells are derived from the hypophysis adjoining the
embryonic cell.
na which has
significance of the phenomenon of APOGAMY (
The physiological ),

recently been accurately investigated in several Angiosperms, has already been

considered (p. 302). Thorough investigation of the embryology has shown that in
these cases we have to do with apogamy and not, as was first assumed, with
parthenogenesis. The reduction in number of the chromosomes (p. 84) is omitted
in the development of the embryo-sac-, all the nuclei of which have the vegetative
number of chromosomes they are vegetative nuclei, not generative, and the egg-
;

cell is apogamous and not parthenogenetic. The fact that the loss of sexuality
is particularly shown in polymorphic genera such as Alchcmilla, Hierncium,
518 BOTANY

Thnlictrum, explained by STRASBUR<;I:K as resulting from the

is

excessive mutation which occurred

lias in time this, together with continuous
;

crossing of the mutations in various directions originating from the sanir spec-jo.
would lead to a considerable weakening of the sexuality, and from this the
apogamous reproduction would follow. On the other hand there are extraordinarily
polymorphic genera such as Rosa and Jliiliiix which have only sexual reproduction.
According to the most recent investigations of OSTENFELD and HI >si N in he genus i.i ;. i

Hieraeium is of special interest, since the formation of the embryo within the <vuh;
may commence in very various ways. In most a tetrad formation accompanied by
a reduction division takes place, but only some of these ovules are found to have a
normal embryo-sac capable of fertilisation as a rule this is displaced by a vege-
;

tative cellwhich develops into an aposporous embryo-sac. In exceptional cases

apogamous embryo-sacs are formed. Thus in ///V/v/c////// apugamy and ap<isp<>r\
are found in addition to the normal mode of development of the embryo as a result
of fertilisation. It is noteworthy that these plants of Hirr/n-him in the course of

Fl(i. :~>OC>. Yonnn eiiiln vo nl

". C.Coty-
FIG. 505. Stages in the development of the emliryo of Capsxlln r, LM'owinx l*>iiit
bvrsa pastoris (A-I>). //.
hypophysis ft. suspc-n^,!
: ,-,,t\-
: ,-.
(Alter HANS I I IN.
ledons p, plumule. (After HANSTEIN, ma^iiili.-ci.
; ) fled.)

generations may, after a larger or smaller number of apogamous or aposporous

embryos have been produced, give rise to descendants which exhibit normal
fertilisation. CLEISTOGAMY ( llb ) may be briefly referred to here, since GOEBKI. has
recently succeeded in reducing it to the same point of view. Cleistogamous flowers
which frequently occur in linpaticns, Lamium, Viola, etc. are to be regarded as
arrested formations, in which, however, the sexual organs attain maturity. They
result in all cases from insufficient nutrition
whether due to want of light or to an
insuflicientsupply of mineral food materials. Since such cleistogamous flowers
regularly set seed, the plants possessing them can succeed even though their
normal chasmogamous flowers are often sterile. Reference may also be made to
the various and often striking provisions which ensure the efficient nutrition of
the embryo and endosperm. While in the simplest cases the endosperm increases
greatly in size and gradually displaces the nucellus, the antipodal cells as lying
next the chalaza are often specially concerned in the nutrition of the embryo-sac.
In such cases they increase in number, and sometimes attain a sixe. Other
large
portions of the embryo-sac itself may grow out as long haustoria which sometimes
emerge from the micropyle and sometimes penetrate into the chalazal region and
the surrounding tissue ; the abundant reserve materials which in some cases
 PHANEROGAMIA 519

(species of Utricularia) are accumulated beforehand, are thus made available for
the nutrition of the macrospore.

12
The Fruit ).
While the formation and structure of the seed
(

presents nothing peculiar to the Angiosperms, the fruit in that group

exhibits a great variety in its development and requires further
consideration here.
Such a variety in the fruit might be anticipated from the range
in structure of the gynaeceum described above. The product of the
individual carpels associated in such apocarpous gyriaecea as those
of the Rosaceae will here be termed PARTIAL FRUITS or FRUITLETS,
while the product of the whole gynaeceum will be spoken of as the
FRUIT. The hollowed-out or projecting floral axis bearing the carpels
may be included in the fruit. In the case of fruits resulting from
syncarpous gynaecea the further development of the wall of the
ovary as the PERICARP has to *be especially considered. The outer-

^ -
n>
FIG. 507. Modes of dehiscence of capsulai- fruits. A, H, capsule of Viola ti-i<-<ili- before and after
the! dehiscence ; C, poricidal capsule of Antii ihinimi iM'jVf (iniiuiiiliid) D, K, pyxidium of
;

Aiiagallis a?vi-n*is before and at'tei dehisceiice.

most, middle, and innermost layers of this are distinguished as

EXOCARP, MESOCARP, and ENDOCARP respectively. According to
the nature of the pericarp the forms of fruit may be classified as
follows :

1. A
fruit with a dry pericarp, which opens when ripe, is termed
a CAPSULE (Fig. 507). When dehiscence takes place by a separation
of the carpels along their lines of union the capsule is SEPTICIDAL ;
when the separate loculi open by means of a longitudinal split, it is
termed LOCULICIDAL, and when definite circumscribed openings are
formed, it is termed PORICIDAL. frequent As special types of
occurrence may be mentioned The FOLLICLE, which
capsule
: is a
developed from a single carpel and opening by separation of the.
ventral suture, e.g. Aconitum napellus (Fig. 594) The LEGUME or :

pod, which differs from the follicle in dehiscing by both ventral

and dorsal sutures, e.g. Cytisus laburnum (Fig. 642) The SILIQUA :

consists oftwo carpels and typically dehisces at maturity by their

separation from a septum lying in the plane of the sutures. The
520 BOTANY PART II

fruit of the Cruciferae, e.g. Cheiranthus Cheiri (Fig. 612), is of this

nature. The PYXIDIUM opens by the removal of a lid, e.g. Anagallis
(Fig. 507).
2. DRY INDEHISCENT FRUITS have a
dry pericarp which does
not open at maturity. Those with a hard pericarp are termed
NUTS. An indehiscent fruit containing one seed, the coat of which

Kit;. "(OS. Dry fruits.

iiideliisci-ut .4, Xut of / Fl<;. OCHt. Schi/nrurp of Hnl.ium
nffidnalis (x ff). B, Achene of Fugopi/rtim esculentum mollvgo. (After DUCHARTRE,
(x 2). (After DUCHARTRE.) x 6.)

isadherent to the leathery pericarp, is known as a CARYOPSIS

(e.g.
Grasses). When the seed is distinct from the pericarp, as in the
Compositae, Polygonaceae, etc., it is termed an ACHENE (Fig. 508).
3. When a
dry fruit, consisting of several carpels, separates at
maturity into its partial fruits without the latter opening, it is
termed a SCHIZOCARP (e.g. Umbelliferae, Malva, Galium, Fig. 509).

Fi<;. 510. Fruit of J'tii/intix c/kekengi, ;. Fruit of Rosa alba, consisting of the fleshy
">11.

consisting of the persistent calyx hollowed axis s', the persistent sepals s, and the

x, surrounding the berry fr, derived carpels fr. The stamens e have witliwii. (After
from the ovary. (After DCCHARTRE. ) DUCHARTRE.)

4. A BERRY is a fruit in which all the layers of the pericarp

become succulent, as in Vaccinium, J^itis, etc. ; when at complete

maturity the succulent tissue dries up a DRY BERRY results (Capsicum).

In a few cases berries may be dehiscent, as in Myristica and some
Cucurbitaceae.
5. In the DRUPE the is differentiated into a succulent
pericarp
 PHANEROGAMIA 521

exocarp and a hard endocarp. Prunus Cerasus (Fig. 627), and

Juglans regia (Fig. 559), are familiar examples.
Fruits which do not originate from the ovary alone are found in Rosa (Fig.
511), in which the succulent receptacle protects the partial fruits and still bears
the calyx on its margin, and in PhysaMs (Fig. 510), the berry of which is enclosed
in the greatly enlarged calyx.
When on the other hand the group of fruits borne on an inflorescence has the
appearance of a single fruit the structure
may be termed a SPURIOUS FRUIT. The Fig
(Ficus) is the best-known example of this,
but similar spurious fruits are especially
frequent in the Urticaceae and Moraceae.
The comparison of a Blackberry which is
the product of a single flower with the
spurious fruit of the Mulberry will show
how closely the two structures may re-
semble one another (Fig. 512).
The above enumeration of the most im- Fit;. 512. .4, Merocarpic fruit <>f Ruims fruti-
cosus consisting of a number of drupes
portant and frequently recurring forms of '' ;

inflorescenc of Mulberry (Moms nigra)

fruit is in no sense exhaustive. It should f
'

bearing a number of small drupes. (After

be added that the various forms are only DUCHARTRE )
to be understood when regarded from an

cecological point of view. The form and other peculiarities of the fruit must
be considered in relation to the distribution of the seeds. In the distribution of
fruits and seeds, as in the transfer of pollen to the stigma, different means of

transport, especially wind, water, and animals, are made use of by plants. (Cf.
the account in the physiological section, p. 318.)

SUB-CLASS I

13
Monocotylae ( )

The Monocotyledons, or Angiosperms which possess a single

cotyledon, are in general habit mostly herbaceous, less frequently
shrubs or trees.
In germination the radicle and hypocotyl of the small embryo
emerge from the seed-coat, while the sheath-like cotyledon usually
remains with its upper end within the seed and absorbs the materials
stored in the endosperm, which is usually well developed. The
growth of the main root is sooner or later arrested and its place
taken by numerous adventitious roots springing from the stem. In
the Grasses these are already present in the embryo within the
seed. Thus a single root system derived by the branching of a
main root, such as the Gymnosperms and Dicotyledons possess, is
wanting throughout the Monocotyledons.
The growing point of the stem remains for a longer or shorter
time enclosed by the sheath of the cotyledon. Later it bears in
522 BOTANY

two-ranked or alternate arrangement the- leaves, which have long

sheaths and continue to grow for a considerable time at their bases.
The growth of the stem is often limited branching is in many cases ;

entirely wanting, and rarely results in the development of a highly

branched shoot-system. The leaves are mostly sessile and parallel-
veined, and of a narrow, elongated, linear, or elliptical shape
(Fig. 513).
Anatomically the Monocotyledons are characterised by their
closed vascular bundles in which no cambium
is
developed ;
these are uniformly scattered iu
the cross section of the stem (cf. Fig. 101).
Secondary thickening is consequently wanting
in Monocotyledons, and in the rare cases in
which it is found results from the formation
at the periphery of the central cylinder of
additional closed bundles embedded in ground-
tissue (cf. p. 144).
The flower in the Monocotyledons is usually
pentacyclic and has two g
whorls constituting the peri-
anth, an androecium of two
whorls, and a gynaeceum of
a single whorl. The typical
number of members in each
whorl is three. The two
whorls of the perianth are
usually similarly formed and
Kit;. ">!;!.- I .i-,-it'o !'/'<' thus constitute a perigone ;. 514. Diagram of u
with
(Fig. 5 1 4). The floral formula
multijliiriiiii piinillrl typical Monocotyle-
venation. (:} nut. six*-.) donous llower.
of such a flower is P 3 + 3,
A 3 + 3, G (3). An increased number of whorls, or of the members
within a whorl, is
only found in the Helobiae.
Iu this and in the apocarpous" gynaeceum of the Helobiae an approach to the
Polycarpicae, among the Dicotyledons, can be recognised, and the attempt has been
made to derive the Monocotyledons from the polycarpic Dicotyledons by means of
this order. This assumption is further supported by the spiral arrangenieht ( !:til ) of
the carpels, and apparently of the stamen* also, in the genus Sa/jittui-iu. This
possible origin of the Monocotyledons deserves more thorough investigation since no
other has yet been indicated, and the Polycarpicae have formed the point of
departure of a number of divergent lines of descent.

Order 1. Helobiae

Water
or marsh plants with actinomorphic flowers. Gynaeceum,
frequently apocarpous carpels in two circles. Seeds, exalbuminou's ;
;

embryo, large. Indehiscent fruits.

 PHANEROGAMIA 523

13b
Alisma Plantayu, Xagittaria sayittifolia and
Family 1. Alismaceae ( ). ,

Butomus ninlifUntus liave long-stalked panicles or umbels, and occur as marsh-

FIG. 515. Floral diagram of

/.Y7//'H'*'i>M'- 1*1 one
rent a.*, Fiii. -'ill'. Suijittm-in *ii<i!ttlf<jlut. a, Flower; /<, fruit attt-r rc-
of tJn> Alisintirrae. (After inoval of some of the carpels. (Magnified ; b, alter Evoun
EICHLER.) and PBANTL.)

plants. The individual flowers have a calyx and a white (in Butomus, reddish)

FIG. r>l~. l'<,t<n,iiaii't:iti initial*. FliiwiTin- .-lioi.i. (J nat. .sj/>>.)

corolla. Androecium, in Butomus A 6 + 3, in Alisma A 6. Gynaeceum apocar-

524 BOTANY PART ij

pous, in JJutomus G
3 + 3, in Alisma G oo SagMaria is mnmccious with
.

flowers tliat by suppression of stamens or carpels are unisexual. Male flowers,

with numerous stamens and sterile carpels female flowers, with staminodes and
;

numerous free carpels inserted on the convex floral- receptacle (Figs. 515, 516).
Leaves in Butcmius, linear, channelled, and triangular in cross-section in Alisma
;

and Sagittal-la, long - stalked, with spoon -'shaped and sagittate leaf -blades
Respectively. -Individuals of both genera growing in deep flowing water have
Jong ribbon shaped leaves, similar to those that appear as a transition type
-

in germination such plants do not flower.

: \

Family 2. Juncaginaceae. Habit grass-like, Triglochin.

Family 3. Potamogetonaceae (Fig. 517). Many species of Fotamoyeton are dis-
tributed over the earth in standing or flowing water. Flowers hermaphrodite,
tetramerous. Gyna'eceum apocarpous. Inflorescence a spike. Leaves usually
submerged, with a long sheath, slit on one side, formed from the axillary stipules.
]'. mitatis, the common Pond-weed, at the time of
flowering has usually only float-
ing leaves, the cylindrical, submerged water-leaves having disappeared by then.
Ruppia maritima and Zanichdlia palustris grow in brackish water. Zostcm
marina. irass- wrack, occurs commonly on all north temperate coasts and is used for
(

stuffing cushions (cf. p. 307).

Family 4. Naiadaceae. The single genus Naias contains a number of fresh-

water plants with diclinous flowers.
Family 5. Hydrocharitaceae. Hydrocharis moi-sim runnr and Stratiotes aloidfs
are floating plants occurring in Britain, which are vegetatively propagated by
runners they pass the winter at the bottom of the water, in some cases as special
;

winter buds (cf. j>. 301), and grow up again in the spring. Flowers dioecious ;

perianth differentiated into calyx and corolla. The male flower has several
trimerous whorls of stamens the female flower possesses staminodes and two
;

trimerous whorls of carpels. Ovary inferior, entomophilous. Vallisncria spiralis

(cf. p. 307). Elodea canadensis, the Canadian water-weed.

Order 2. Glumiflorse

This order consists entirely of annual or perennial plants of grass-

like habit. A woody stem only appears in the genus Bambnsa. The
association inmore or less complex inflorescences of numerous flowers,
which lack a proper perianth but are enclosed by scaly bracts (glumes),
is a common character of the order. The perianth is either completely
wanting or reduced to a series of scales or bristles. The inner whorl
of stamens is also wanting. The superior ovary is always unilocular
and contains only one ovule ; it is formed of three, two, or of a single
carpel. The large size and feathery and papillose form of the stigmas
stand in relation to the wind pollination. Fruits indehiscent.
Family 1. Cyperaceae. The Sedges are characterised by their
triangular stems, which are usually neither swollen at the nodes nor
hollow, and by their closed leaf-sheaths. The flowers are unisexual
and usually moncecious or are hermaphrodite ; ovary formed of two
or \ three carpels with an erect, basal, anatropous ovule. Pericarp
not coherent with the seed-coat ; embryo small surrounded by the
endosperm.
 PHANEROGAMIA 525

Scirpus, and Eriophorum have hermaphrodite flowers.

The genera Cyperus,
Fig. 518 represents a plant of Scirpus setaceus, which, is an annual, in flower.
Leaves rigid, channelled above. Fertile shoots with the uppermost internode

elongated. Spikes 1-3, terminal enclosed hy imbricating bracts and displaced

;

to one side by the subtending bract the line of which continues that of
the stem. Only the large, lowermost bracts are sterile, the others have each
a naked hermaphrodite flower in their axils. Other species have the perianth

FIG. 518. Scirpus setaceus. 1, plant in

flower ; upper portion of a flowering
2, I'h-. .">!!'. Eriaphorum angnstifullmn, 1, Inflores-
shoot ; 3, single flower 4, the same from
; cence ; 2, a single spikelet ; 3, single flower ;

behind ; 5, the same without the bract ; it, flower with bract removed ; 5, fruit. (After
6, fruit.(After HOFFMANN. 1, nat. HOFFMANN ; 1, about nat. size ; the others x
size, the others x -'-('>. ) 3-5.)

represented by bristles. The Cotton-grass (Eriophorum anyustifolium], which

when flowering is bears at the summit of its fertile shoots
inconspicuous,
3-7, long-stalked, erect spikelets with numerous, imbricate bracts. Around the base
of each flower are numerous hairs, which
ar^ concealed by the projecting stamens
and style. When the plant is in fruit the hairs, which have become about 3 cm.

long, project freely from between the bracts and constitute a valua-ble means of
dispersal for the fruits. The white colour of the hairs makes the now pendulous
spikelets of the Cotton-grass a conspicuous feature of peat-moor vegetation
(Fig. 519). Cyperus papyrus, in Egypt and Sicily.
The genus Carex is for the most part monoecious, and its flowers are naked and
526 HOT A \ Y PART II

unisexual. Male spikes simple ;

in the- axil of each bract la a male flower formed
of three stamens. The female spikes hear in the axil of each hraci secondary ,i

shoot; the axis of this is included in the tubular subtending bract (utriculus)
together with the pistil (formed of 2 or 3
Q ttfr. O carpels) which is borne in the axil of the
latter (Fig. 520).

Family 2. Gramineae. The

stems of the true Grasses are cylin-
drical, and have hollow inter nodes
(exceptions Maize and Sugar-cane) ;
the nodes are swollen ; the leaves
are two- ranked and their sheath is
usually split and thickened at the
node. At the junction of the sheath
and leaf-blade, a membranous struc-
ture (the ligule) projects (cf. Fig.
of a male ttowei
i

36). The flowers of the Gramineae

of OWN: :, of a female flower with three j
s, i,,,,,s ;<-.,,. -a female ,., with two
are grouped in spicate, racemose, or
* ;D, diagram of a female flower paniculate inflorescences, which are
;S, diagram of the hermaphro- always composed of partial inflores-
clite spikelet of A'///"" :<'. secondary axis : , i_ i *. TT n
,rtr, utricuitw or bract of the secondary cences, the spikelcts. Usually each
axis. (After EICHI.ER.) SPIKELET bears several flowers. At
the base of the spikelet there are
usually a pair of sterile bracts (GLUMAE); sometimes there is only one,
or 3-4, glumes. Continuing the two ranked
-

arrangement of the glumes, come the fertile sub-

tending bracts (PALEA INFERIOR) in the axil of each
of which stands a flower. The subtending bracts
are often awned, i.e. they bear terminally or
springing from the dorsal surface a stiff bristle
with backwardly directed hairs (the AWN). The y
, bracteole of each flower is
represented by another scale-
like bract, the PALEA SUPER- 4
IOR. Above this come two
small scales, the LODICUL/E,
the distension of which assists ;. :"r_'-J. --Diagrammatic re-
in opening the flower. Lastly jirest'iitJitioii of a (irass

the axis bears the androecium sjiikelet. r/. The glumes :

;>, and /M, tlir iiifci'ioi

iiia -ram consisting of a whorl of three and siijxTior jialea ; .

oftheGramin<>ae04wn) stamens, and the ovary com- Icxlicilles : /;, tlovver. Tlie

Thr absent members are axial parts are re|n.--

poge(] Q f Qne
x.
carpe l an(J s.'iit-jd a-; elongated.
represented l>v r e ^i -n
bearing two feathery papillose
stigmas. The ovary contains ;ui anatropous, or slightly campy lo-
tropous ovule (cf. Figs. 521-523).
The flowers do not always show such extreme reduction thus the flower of :
 PHANEROGAM A I 527

Rice (Fig. 526) has a complete androecium that of the Bambuseae is similar and ;

also has three lodicules, and in Strcptochfrta there is a normal monocotyledonous

type of flower with all five whorls of members present (cf. the diagram in Fig. 514).
The lodicules can on this evidence be
regarded as corresponding to the inner
whorl of the perianth. Possibly the
superior palea represents two coherent
leaves of the outer whorl, but proof of
this is at present wanting.

The length of the staminal

filaments, attachment a
their
littlebelow the middle of the
anthers, and the extended surface
offered by the feathery stigmas
are of value for wind-pollination
(Fig. 523). The fruit of the

KKI. .V24. Part of median longitudinal section of a

grain of Wheat, showing emliryo and scutellum
Fie;. .123. fr'i'ftiK-H i luli', i'. A. Spikelct (compare Fig. (so); tv, vascular bundle of scntellnm ve, its ;

.VJ2)with two open Mowers below which the two sterile columnar epithelium ; /', its ligule ; e, sheath-
glumes are seen(x 3); B, Flower; tin two lodicules ing part of the cotyledon pv, vegetal
1

arc in front, the superior ]>alea behind the ovary bears ; of stem hji, hypocotyl I, epiblast r, radicle
: ; ; :

two feathery stigmas (x 12); C, a single lodicule <!, root-sheath m, micropyle p, funieulus vp,
; ; ;

(X 12); D, ovary seen from the side with the >talk its vascular bundle ; /, lateral wall of groove.
of one of the removed stigmas ( x 12).
-
(X 14.)

Grasses is termed a caryopsis ;

in it the pericarp and seed-coat are
intimately united. The embryo lies in contact with the endosprrm
by means of its cotyledon ;
this forms the SCUTELLUM and in germina-
tion serves as an absorbent organ by means of which the reserve
materials in the endosperm are taken up by the seedling (Fig. 524).
The most important economic plants belonging to this order are the Cereals
528 BOTANY PART II

(Fig. 525). Wheat, Triticum. Spikelets single, with two or more flowers glumes ;

broadly ovate. KOERNICKE distinguishes as spccirs <>t' Wheat, 1. Tr. vulgare, with
a number of sub-species; 2. Tr. polonicum 3. Tr. innnococctini. Rye, Secale
:

FIG. 525. Cereals. A, Rye, Secalfcftfulf : I',, .S]H-k. Triticmii s^'lti, ; r. T \vn-i ankf! Barli-v,
Hordeum vulgare, tlixtii-hinn ; 1>. \Vlit-at. Ti-itiriim m/iim-i:

ccrcale; Spikelets single, 2-flowert'il glumes acute. Barley,- I/of/' nm. nilgare;
;

Spikelets one-Howered, in groiijis of three in the Mib-species H. hcxastichum

;

and H. letrastichuin all the rows of spikelets are fertile, in //. 'Hxt!<-hum only the
middle row. Oat, Avena saliva. Maize, Zea mais. The above are all cultivated
in temperate climates, the Maize, largely in America, the others also in Western
 PHANEROGAMIA 529

Asia, and the south-east of Europe. In the wild state only Triticum aegilopodioidcs
(from which Tr. monococcum is derived), Secale montanum, and Hordeum spontaneum
(allied to H. distichum) are known. In these wild forms, the spikelets fall from
thi' rhiichis at maturity, a character that would be unsuitable in cultivated forms.
The most important tropical food-plant of the order is Rice, Oryza saliva (Fig.

B
FIG. 5-26. Onizn sativa. Panicle (J nat.
size), mid a single spikelet (enlarged.) Loliiim temulentum.
OmciAL. Pouosoua.

526), which is largely cultivated to the limits of the warmer temperate regions,
and, when sufficient moisture an enormous harvest. In Africa,
is available, yields
several varieties of Millet, Andropogon Sorghum, are cultivated, and it forms the
most important cereal for that continent. Panicum miliaceum and P. italicum,
of Asiatic origin, are still cultivated, though to a diminished extent, in the
Mediterranean region. The Sugar-cane, Saccharum qfficinarum, is another
important food -plant; it is a perennial, growing more than six feet high, and
2 M
530 BOTANY I'AKT n

occurs in tropical Asia. The Sugar-cane is cultivated in all tropical countries,

and cane-sugar is obtained from the sap expressed from the solid stem.
Among the most important of our meadow - grasses may be mentioned
Agrostis alba, Alopecurus pratensis, Ant/iii.'-in/f/n/iti odoratum, Arrhenatherum
elatius,Arena flavescens, A. pubescens, Briza media, Dactylis glomerata, Holcus
lanatus, Lolium perenne, Phleum pratense, Poa pratensis, and species of Aira, '
Bromus, Calamayrostis, Festuca, Melica, etc. The tropical species of Bambusa,
which grow to the height of trees, are utilised in many ways from the stems are ;

constructed houses, walls, flooring, ladders, bridges, cordage, water -

vessels,

cooking utensils, water-pipes, etc., and the plant is indispensable in the countries
in which it occurs.
POISONOUS. Lolium temuletttum (Fig. 527) and L. mn,l\iin have poisonous
fruits in relation to which fungal hyphse occur these plants are both annuals and
;

can be distinguished by the absence of sterile shoots from the common Lolium
perenne and L. multiflorum.
OFFICIAL. AMYLUM (starch) is obtained from Triticum sativum, Oryza saliva,
etc. ;
SaccJiarum officinarum provides SACCHAurM.

Order 3. Spadieiflorae

The common character of this order is afforded by the peculiar

inflorescence ;
this is a spikewith a thick, swollen, often fleshy axis
and is termed a spadix. The flowers are mostly diclinous,
monoscious or more rarely dioacious.

Family 1. Typhaceae. Marsh plants, with long, linear leaves and long-
stalked spikes, which bear a large number of flowers, the male above, the female
lower down. Perianth wanting.
Family 2.
Sparganiaceae. Connected with the preceding family. Spikes
spherical. Flowers with a perigone, but otherwise like the Typhaceae.
Family 3. Pandanaceae. Trees of peculiar appearance, supported by prop-
roots, or climbing, shrubby plants all belong to the tropical countries around the
;

Indian Ocean and to the Pacific islands. Leaves elongated, .spiny, channelled
above, arranged without bare internodes in three ranks on the axis. Inflorescences,
<Jor?, are terminal spikes in the axils of sheathing bracts. Flowers without
perianth, Pandanus (cf. Fig. 531 in front of the Palms), Frcycinetia.

Family 4. Palmae. The Palms are an exclusively tropical and

sub-tropical family, the members of which mostly attain the size of
trees. Their slender stem is simple and usually of uniform diameter
throughout ; only the African species of Hyphaene have branched
stems. Other forms show evident growth in thickness towards the
base ; depends on enlargement of the elements already
this either
present, or resembles the growth in thickness of arborescent Liliaceae.
The leaves, which are often of gigantic size, form a terminal crown.
They are either pinnately or palmately divided, the division
coming
about by the death of definite portions of tissue in the young leaf in
Fin. KS&^COOM HVCtftra. Intlori-si-.-no- of the Cucu-nut 1'ulin. (Greatly reilm-eil.)

531 2 M 1
532 BOTANY PART II

the bud. The inflorescence is in some cases terminal (Metroxylon), and

the individual perishes with the development of the fruits. More
often the inflorescences are axillary. When young, they are enclosed
by a massive resistant sheath, the spathe this bursts open and permits ;

of the unfolding of the simple, or more usually branched, inflorescence.

The individual flowers are as a rule unisexual and constructed on the ordinary
monocotyledonous type ;
P 3 + 3, A 3 + 3, in the male flowers, and P 3 + 3, G (3),
in the female flowers. In Cocoa their distribution is moncecious. Fig. 528 repre-
>riits the inflorescence of (,'</<</.< /(//<//'/". still partly enclosed by the spathe. The
male flowers are crowded on the
terminal brandies of the inflores-
cence, while the female flowers
are considerably larger and stand
singly lower down. The ovary,
which is here composed of three
united carpels, becomes, as a
rule, unilocular in the fruit,

i;.530. Coco-nut after partial re-

moval of the fibrous exocarp. (After
Fin. .")-".. ClK-l.l-Iillts (III till' Stem. WAGNER, reduced.)

since only one carpel develops further. The ripe fruits (Fig. 529) are borne in
small numbers on each inflorescence. Each consists of a coarse, fibrous exocarp,
which contributes to the buoyancy of the fruit in water, and thus leads to the
wide distribution of this palm on tropical coasts, and a hard endocarp on which
the sutures of the three carpels can be plainly distinguished. At the base of each
carpel a germinal pore is present in the endocarp (Fig. 530), but only the one in
relation to which the embryo lies, remains permeable. The endosperm forms a
thick layer within the endocarp it is rich in fatty substances and produces the
;

COPRA of commerce. The space within the endosperm is partially filled with fluid,
the "milk" of the coco-nut, which is possibly of service in germination. The
embryo on germination develops a massive absorbent organ which grows into
the cavity of the fruit and serves to absorb the reserve materials. Fig. 531 shows
the general habit of Coco-nut palms.
Differences are, however, found within'the order. In Arcca catechu, the fruit
developed from a similar ovary to that of Cocos is a berry, the exocarp becoming
partly fibrous and partly succulent. The white endosperm is here of stony
SECT. II PHANEROGAMIA 533

consistence, cellulose being stored as a reserve material the endosperm is

;

ruminated, i.e. the dark seed-coat grows into it at many points and gives it a
veined appearance. The fruit of the Date Palm (Phoenix dactylifera) is also a
berry, but this arises from one of the carpels of the apocarpous gynaeceum, the
other two not developing. In contrast to the other genera mentioned, Phoenix

irt I'alins at Hiln. Hawaii : 1'niiiliiiuix ill front of the Palni.s

is dioecious. Other important economic plants among the Palms are Elaeis
yuineensis, the African Oil Palm, species of Calamus which yield Malacca Cane,
and species of Afttroxylon, from which Sago is obtained the two latter are found
;

in the Asiatic-Australian region of the tropics. Phytelephas macrocarpa, an

American Palm which does not form a trunk, yields vegetable ivory (the hard
endosperm). Several species yield a flow of sugary sap on cutting off the
inflorescence, and this is sometimes fermented to make Palm-wine, sometimes used
as a source of cane-sugar (Arenya mccharifera).
2 M 2
534 BOTA N Y PAKT II

Family 5. Araceae. The Araceae are mostly herbs or shrubs ;

they take a conspicuous place as root-climbers in the damp tropical

forests. The leaves of some species ((>.<(. Motutera) have the large
lamina incised or perforated this comes about by the death of
;

Flmvrrinx ]>l:nit. Sin^li- MU\V.T> sr.-ii from iilMivi- :ui<l t'nmi

tin- side, (i n:it. >,ixi\)

definitely limited areas and is comparable to the method by which

the leaves of Palms become compound. Flowers are greatly reduced,
usually diclinous, borne on a swollen, fleshy axis a spathe, often of ;

bright colour and serving to render the inflorescence conspicuous,

 PHANEROGAMIA 535

is present, at the base of the spike (e.g. Anthurium scherzerianum,

FK;. 533. Artamacvlatwn(jfra.t. size), [ntioreacenee and fruits ('j nat. size). Poisoxovs.

Richanlia aethiopica, both of which are commonly cultivated). Fruit

usually a berry.
2 M 3
536 BOTANY r.M.r ii

Acorus calamus has, in the course of the last two or three centuries, spread to
this country from the East. It has complete, hermaphrodite flowers; ovary tri-
locular. The short spadix is terminal, but is displaced to one side by the sp.it he.
which resembles the foliage leaves (,Fig. 532).
POISONOUS. Many Araceae are poisonous. Calla palustris in peaty swamp-.
Arum maculatum (Fig. 533), a perennial herb with tuberous rhizome, common in
woods. It develops a number of stalked, ha.-taie leaves, the brown spots on which
give the plant its specific name. The flowers are monoecious, without perianth ;

the female flowers stand at the base of the spadix and the male a short distance
above them. Above the latter come a number of sterile flowers with downwardly
directed, hair-like j mints, which stand at the level of the constricted i>ortion of the
spathe this is widely open above.
;
These hairs allow insects, attracted by the
peculiar scent or seeking warmth (cf. p. 244), to creep into the lower expanded
portion of the spathe, but prevent their return until the female flowers have been
pollinated from another individual. When this is accomplished the hairs wither
and the anthers open. The escaping insects, now dusted with pollen, may enter
other inflorescences and pollinate the flowers.
Family 6. Lemnaceae. The Duck-weeds of our ponds and streams have naked
monoecious flowers borne in depressions of the body of the plant. The flowers are
surrounded by a small spathe. Lemna, Wolffm.

Order 4. Enantioblastae
A small order. Ovules orthotropous ; embryo at summit of endosperm at

opposite end from the hilum.

Family Conimelinaceae. Peri :
anth developed as calyx and corolla.
The hairs of the stamens all', rd
well-known objects for the study
of movements of protoplasm and
nuclear divisions. Fruit a capsule.
Comineliiia, Ti'in

I-li.. .
liinij'i-iH-iir/iii*. a, 1'jirt lit' an
Flo. 534. Diagram nt'u typical I.iline.'<>ii> illtldl-escelice I single fll IWcr (h) ;ill'l .H.MI
-

flower. aeceiun (<) iim^nilifil.

Order 5. Liliiflorae

Flowers actinomorphic, composed of five whorls, \\ith superior or

inferior ovary. Both whorls of the perianth developed alike. Only
 PHANEROGAM A I 537

ii.. Viii. i'l.-l,ii-in initinnnalf. (i nut. size.) PoiSOXOUb ; OwciAL.

538 BOTANY

in the Iridaceae is one whorl of the androecium suppressed. The

gynaeceum varies in position, but it is always formed of three carpels

and in most cases encloses a tri-

locular ovary.
The plants of this order show
a wide range of habit. Besides
herbaceous forms with bulbs or
corms or subterranean rhizomes,
climbing shrubs, succulent and
armed, rosette - forming plants of
large size and larger or smaller
trees occur ;
the latter are re-
stricted to tropical and sub-tropical
regions.
Family 1. Juncaceae. Plants of

grass* like habit. Flower ot complete

Liliaceous type ;
with
scaly perianth.
Wind -pollination. Pollen -grains united
in tetrads. Ovary superior, uni- or tri-
locular, bearing three, long, papillose
stigmas. Endosperm floury. Fruit a

oapaule.
Numerous species of JUHCHX (Rush)
occur in our marshy ground the
flora, in :

leaves are cylindrical and have large in-

tercellular spaces. The clusters of small,
aiiemophilous flowers (Fig. 535) are borne
on the end of a shoot but are often dis-
placed to the side by the bract which
continues the line of the axis. Fruits
with many seedsi Luzutti, with flat
leaves and three-seeded fruits, one of the
earliest spring-flowering plants.

Family 2. Liliaceae. Typical

flower, with coloured, conspicuous
perianth.Ovary superior. Endo-
sperm horny or fleshy. Divided
into three sub-families according to
FIG. 537. i.'iili'hicum /</,,)"/. Longitud- the construction of the fruit.
inal section through conn of a flowering
(a) Capsule with three styles ;
tin-

plant. Explanation in t'-xt. (After SACHS.)

OffTCUL.
dehiscence septicidal, i.e. fruit splits
through the septa. M<>l(mt)ti<'<i<-.

(6) Capsule with one style dehiscence loculicidal, ;

i.e. fruit splits

opening the chambers of the ovary. Lilieae.

(c) Fruit a berry. Asparageae.

Colcli ii-iiin ini/ttnntfi/f, the Autumn Crocus (Figs. 536, f>:j7), is a perennial herb
growing in meadows. If a plant is examined in autumn at the time of (lowering.
 PHANEKOGAMIA 539

the conn (k), to the base of which

attached the lateral shoot bearing the flowers,
is
will be seen to be enclosed in a (Fig. 537 h). Roots spring from
brown envelope
the base of this lateral shoot (/), while those of the old corm (w) are in a withered
condition. At the summit of the corm the remains or scar of a dead aerial shoot
will be seen (st). The lateral flowering shoot bears at its base three sheathing
leaves (s', s," I') not separated by elongated internodes. In the axil of the third of

f
l-'ic:. .Mis. u-c, Ornithngulu in umbel/alum : n. entire' plant (reduced); '/, Mower
(nat. size); c, flower,
part of ]>rij;oiie and amlroecium removed >/, fruit e, ; ; fruit in transverse section, f-g, t'nli-lii-
ii ni iint" in n'lJi' : /. fruit in transverse section g, section through seed showing endosperm
;
(<).

(<-;/ nmxnitied.)

(I') is a bud (k")

these which will form the flowering shoot of the next season this ;

third leaf will develop a lamina and appear above ground in spring as the first
foliage leaf. A thickened internode (k') separates it from the two leaves (I")
situated higher up on the axis. Above them come the flowers (b, b'), which may
be four in number, and stand in the axils of reduced leaves by the arrest of the ;

growing point of the axis the flowers appear to be terminal. In spring the reserve
materials from the tuber (k) are absorbed and the old tuber is pushed aside by the
swollen internode (k"), which in its turn enlarges to form a new corm. The three
ii;. .'.39.
Urgineamari-
tima (about -fn nat.
size.) OFFICIAL. (After FIG. 540. Aloe socot i ii:n. Plant in flower (greatly reduced); tip of leaf,
BERO and SCHMIDT.) single flower, and cross-section of ovary. (} nat. size.) OFFICIAL.
540
SECT. II THANEROGAMIA 541

foliage leaves expand their long, channelled, dark green lamina above the*soil ;
their sheathing portions closely surround the axis. The latter bears the fruits,
which contain numerous, spherical, black seeds these are liberated by the
;

dehiscence of the capsule at the sutures. The aerial shoot then dies, the leaf-
sheaths surrounding the new corm, which bears a lateral bud (") in the axil of its
first foliage leaf this bud was already evident in the preceding season.
;
Veratrum
album is a conspicuous herb with a rosette of large, elliptical, longitudinally folded
leaves. The growth of the main axis is terminated by an inflorescence, which is a
panicle more than a metre in height the leaves borne on it have long sheaths
;

and diminish in size from below upwards. The greenish -white flowers are

FKJ. 541. Dracaena draco.The dragon tree of Lacuna in tht- Canary Islands.
(From CHUN, Aus den Tiefen des Weltmeeres.)

polygamous, i.e. partly hermaphrodite and partly unisexual the ovary bears three
;

widely-spreading styles.
Such popular flowers as Tulipa (Fig. 25), Hyacinthus, Lilium, Muscari, and
Scilla, and vegetables as Allium, together with Urginea (Fig. 539), which occurs
in the Mediterranean region, belong, on the other hand, to the Lilieae. Ornitho-
galum umbcllatum (Fig. 538) will serve as an example of this group. In autumn
the plant consists of a bulb, each of the fleshy scales of which has a scar at the
upper end numerous roots spring from the base of the bulb. The bulb-scales are
;

more or less grown together. In the axil of the innermost scale is the stalk of the
spent inflorescence together with a young bud bearing a number of leaves. Each
of these leaves is provided with an embryonic lamina, while the continuation of
the shoot is the embryonic inflorescence. In spring the leaves grow into long,
linear structures, and, together with the inflorescence, appear above ground. The
inflorescence is sparingly branched the white flowers have a trilocular ovary
;
542 BOTANY PAIIT II

bearing a common style. Tin- upper parts of the leaves wither while the li:i>;il

KII.. m.l'<ii-isiiii>i'li-;f<>lia.
(1 nat. size.) /'o/>v,..,,,/ i

portions become swollen and fleshy and stored with re.-i.-rvc materials ; the g
 PHANEROGAMIA 543

the upper part of each scale marks the place of separation of the leaf-blade. The
annual course of development is essentially similar in other bulbous plants. The
the cold or, in the
vegetative period is restricted to a few months, while during
numerous bulbous plants of warm-temperate climates, the dry seasons, the bulb is
protected by its subterranean situation. Aloe, a genus of African plants con-
taining many species (Fig. 540) may be mentioned as an example of an arborescent
member of the Lilieae.
Dracaena (Fig. 541), an arborescent form which attains
a great age and a
characteristic appearance, together with the similar genera, Gordyline and Yucca,

belong to the Asparageae. To the same sub-family belongs Smilax (Sarsaparilla),

a shrubby plant of warmer countries, climbing by the help of tendril-like
emergences at the base of the petioles. Other examples are Asparagus with
bunches of phylloclades (p. 25) in place of leaves, Convallaria (Fig. 34),
Maianthemum, Polygonatum, and Paris quadrifolia (Fig. 542) the latter bears;

whorls of four leaves, sometimes 3-6 leaves. All these plants have creeping rhizomes
bearing scale-leaves either the apex of this rhizome grows annually into the
;

erect shoot bearing the foliage leaves and inflorescences, while the growth of the
rhizome is continued by a lateral branch (Polygonatum, Fig. 23), or the rhizome
continues its subterranean growth, the leafy shoots being developed from
axillary buds (Paris).
POISONOUS. Numerous Liliaceae are more or less poisonous, e.g. Lily of the

Valley, Tulip, Fritillaria, Colchicum, Vcratrum, Paris.

OFFICIAL. Colchicum autumnale, seeds and corm Schoenocaulon (Sabadilla)
;

a bulbous plant with grass-like foliage from mountains of Central

officinalis,
America and Venezuela, seeds yield VERATRINUM. Aloe vera, A. chinensis, A. perryi
and other species yield ALOES BARBADENSIS and ALOES SOCOTKINA. Urginea scillae
yields SQUILL. Smilax ornata, etc., yield SARSAPARILLA.
Family 3. Amaryllidaceae. Distinguished from Liliaceae by the inferior
ovary. Lcucojum (Fig. 543), the Snowdrop (Galanthus) and Narcissus resemble
the bulbous Liliaceae in habit. Agave, large plants with succulent leaves from
the warmer regions of America, provide fibres. A. mexicana provides the national
drink of Mexico (pulque), obtained by fermenting the sap that flows on cutting
off the inflorescence. Species of Agave are naturalised in the Mediterranean region
but none are native there.

Family4. Iridaceae. Distinguished from Liliaceae by their

inferior ovary and by the suppression of the inner whorl of the
androecium (Fig. 544). The two whorls of the perianth are not
always similar. Anthers extrorse. The leaves of the Iridaceae are
always sessile the underground portion is a tuberous or elongated
;

rhizome, rarely a bulb. Capsule loculicidal.

Crocus satimis, Saffron (Fig. 545) is a plant which has long been cultivated in
the East it has a tuberous rhizome and narrow, grass-like leaves. The flowers
;

are sterile unless pollinated with pollen of the wild form. The large
stigmas
furnish Saffron. Other species are cultivated as ornamental plants.
Iris, leaves overlapping in two ranks. The leaf-sheath surrounds the thick
fleshy rhizome, while the sword-shaped blade stands erect and has its two
surfaces alike (Fig. 546). Outer perianth segments bent downwards, inner erect.
The three anthers are roofed over by the three, leaf-like styles. In Gladiolus the
flowers are dorsiventral and the dissimilarity of the perianth leaves is more
marked.
544 BOTANY I'AKT II

OFFICIAL. Crocus sutivua yields SAFFKOX.

Family ;">. Climbing shrubs with diclinous flowers, sagittate or
Dioscoreaceae.
cordate, frequently net-veined leaves and large, tuberous rhi/oiiirs. Mostly tropical.

r
Km. . i43. Leticoji'm nt'ftinnn.

a, Inflorescence (redact-d) ;

/', jj.MiHeeeiiin anil andror-

ciuni (nat. '.size).

KH;. f)44. Floral rlia^-;iiii c.l *ttii-n.->. Style with tripartite stigma.
the Iriilaceae (// /'.<). OFFICIAL. (After HAILLOX.)

IHoscorea batatas, Yam (China) ; Testudinaria elephantipes is often cultivated.

Tamus communis, the Bryony, is British.
Family 6. Bromeliaceae. Mostly epipliytcs : tlowt-rs hermaphrodite, position
 '

FK.. ."ili'i. -/ (} " ; 't- MXP.)

545 2 N
546 BOTANY

of ovary variable. Limited to tropical and sub-tropical parts of America. The

leaves are in rosettes and are typically xerophytic in the forms which grow in
;

the soil they are spiny. Ananassa saliva is cultivated and has spread in the
wild state in the tropics ;
its inflorescence forms the Pineapple.

Order 6. Seitamineae

Tropical plants, sometimes of large size, in a few cases arborescent.

Flowers dorsiventral or asymmetrical. Perianth differentiated into
calyx and corolla. Androecium greatly reduced ; some of the
stamens represented by staminodes, and resembling the segments
of the corolla. Ovary in-

ferior, trilocular. Seeds

with perisperm.

Family 1. Musaceae. The

Banana (Musa) is one of the
most important plants of all
tropical countries. The appar-
ent, erect stem is formed of the
closely overlapping, sheathing
bases of the large leaves. In-
florescence, terminal. Fruit, a
berry. M. textilis yields Manilla
Hemp.
Fio. 547. Floral diagram of Zingiberaceae (after EICHLER).
b, Bract v, bracteole ; k, calyx pi ., segments of corolla;
; ; _, ..

ssti,sst.>, staminodes
of the outer whorl of the androecium; .family 2. Zingiber-
* the
suppressed stamen of this whorl ; ft, the singlr aC6ae. Flowers in spikes,
fertilestamen; I, petaloid staminodes of the inner
w hi ch : ,.
.- p ,

whorl of the androecium forming what is known as the

labeiium. semble capitula. Flower
dorsiventral. Calyx incon-
spicuous, tubular. Corolla with three lobes. Outer whorl of the
androecium wanting or represented by two lateral staminodes
is

(Fig. 547, sst v sst z ). Only the posterior stamen of the inner whorl
(st) is fertile; the two others are joined to form the brightly
coloured, petaloid labeiium (/). The style lies in the tubular groove
between the two thecae of the stamen. Fruit a capsule. Most plants
of the family belong to tropical Asia.

Zingiber officincde, the Ginger, is an ancient cultivated plant of Southern Asia,

now cultivated throughout the tropics (Fig. 548). The flattened branched
rhizome is in contact with the soil by its narrow side. Leaves, two-ranked ;

main shoot continued by the growth of axillary buds of the lower surface. The
leafy shoots, in spite of their length, are composed of the sheaths of the large,
simple, entire leaves, the axis remaining extremely short. Bracts large and,
especially at their margins, brightly coloured. Flowers, bright yellow, with a
conspicuous, violet and spotted labeiium. Elettaria Cardamomum and Curcuma
 PHANEROGAMIA 547

have the stalks bearing their inflorescences .similarly composed of leaf-sheaths.

sl/jiitiia and Hcdychium, the latter of which is often cultivated, have on the
other hand normal, leafy shoots bearing the terminal inflorescence.
OFFICIAL. Zingiber ojficiiutle, rhizome yields GINGER. Elettaria Cardamomum
yields CAKHAMO.M SEEDS.

Kic. ."j48. Zii i>ffi<:lnnlt. (\ nat. si/t\ After BERI; ami SCHMIDT.)

Family '3. Cannaceae. Large-leaved herbs often in cultivation. ;

Flowers
asymmetrical (Fig. 549). Only one half stamen fertile (i.e. anther with only one
theca) the other half being petaloid.
Family 4. Marantaceae. Large-leaved herbs. Leaves with pulvinus at
junction of stalk and lamina. Stamen as in preceding order. Arrowroot is
obtained from Maranta arundiiiacca.
548 BOTANY

Order 7. Gynandrae
Family Orehidaceae. Perennial, herbaceous plants growing as
epiphytes or in the ground, with hermaphrodite, zygomorphic flowers.
Perianth petaloid, the pos-
terior segment of the inner
whorl developed as a lip or
labellum, which frequently
"
bears a spur. (The label-
"
lum of the Scitamineae
being formed of two stam-
inodes, is entirely different
morphologically.) Androe-
cium formed of the three
anterior stamens only ; the
middle stamen, belonging to
the outer whorl, is fertile ;

FIG. 549. Flower of Cunnu iridifloru. //Ovary ;

k, calyx ; c, corolla I, labellum ; stj-s, the

other staminodiu a, ; fertile stamen g style. Flo. 550. Orchidaeeae. Floral
(J nat size.) diagram (0/r/u.--).

the other two are represented by staminodes.

Cypripediwn has
these two lateral stamens of the inner whorl fertile.
Gynaeceum

FK;. 551. Orchis inHltitriK. Longitudinal Fl<;. 552. Koot-.syste.ni of Urchin latif<>li. 1,,
section passing through the old and Base of stem ; s, cataphyllnry leaf; ', old,
new tubers. (After LCEHSSEN.) I", young tubers /, bud 1

roots.
; .
; r,

formed of three carpels, syncarpous ; ovary inferior, unilocular.

Fruit, a Seeds
capsule. extremely numerous, borne on parietal
 PHANEROGAMIA 549

placentas (Fig. 550). The fertile stamen is adherent to the style

and forms with it the COLUMN or GYNOSTEMIUM this projects more or ;

less in the centre of the flower. The labellum, which serves as an

alighting place for visiting insects, becomes anterior either by the
torsion of the whole flower through 180 (cf. Figs. 550 and 553) or
by the flower being bent backwards.
This order of 6000-10,000 species is primarily divided into Monandrae and
n,

I-'H;. .VVt. o/r/iiy i,iii;tnriri. a. Flower: ".

bract ; li, ovary ; <, the outer, and d, the

two anterior inner perigone leaves >. :

labellum with the spur/; g, gynostemium.

b, Flower after removal of all of the peri-
gone leaves with exception of the upper
part of the labellum h, stigmai; I, rostel-
:

him ; fc, tooth-like prolongation of the

rostellum ; in, anther ; u, connective ; o, Flo. 554. Vanilla planifnHa (reduced. After BERG
pollininm ; </, viscid disc ; p, staminodium. and SCHMIDT from KNCILER and PRANTL.) A,
;

c, A pollinium :
r, caudicle ; ,, pollen. Labellum and gynostemium li, gynostemium
;

d, Ovary in transverse section. (After from the side C, summit of the gynostemium
;

BERd and SCHMIDT.) from in front Z>, anther K, seed. (Magnified.)

; ;

Diaiidrae in which one stamen of the outer whorl or two stamens of the inner whorl
are fertile respectively.
EXAMPLES OF BRITISH GENERA 1. Monandrae Orchis, Ophrys, Gymnadenia : :

with tubers Epipactis, Cephalanthera, Listcra with branched rhizomes Neottia,

-, ;

the Bird's-nest Orchid, Epipogon, Corallorhiza, saprophytie and almost destitute

of chlorophyll. 2. Diandrae Cypripedium, Ladies' Slipper. :

Orchis militaris which is represented in Fig. 555 will serve as an example for
more detailed consideration. At the period of flowering a pair of fleshy tubers
will be found at the base of the plant, both of which are covered with root hairs.
The large or brown tuber of more spongy texture continues above into the stem
550 BOTANY

which terminates in tin- pyra-

midal inflorescence this axis ;

is surrounded at the base ;i

liy

pair of scale leaves and the

-

sheaths of the 2-4 elongated,

elliptical foliage leaves. The
smaller tuber is of tinner con-
sistence and of a white colour :

it bear.-., as is shown ill the

longitudinal section (Fig. 551),
a lnid on its summit which
already shows a pair of scale
leaves. This tuber has arisen
as an axillary bud in relation
to one of the first scale-leave-.
of the plant, and with its
tuberous, swollen, first root has
lirnken through the subtend-

ing scale-leaf (Fig. 555). It

is destined to replace the
parent plant in the succeeding
season. The anatomical and
morphological differences be
tween the tuber and the ordi-
nary roots of the plant may be
explained by the consideration
that the tuber arises from
several coherent roots. The
existence in other species _ ; 1

552) of palmately branched

tuliersincrcases the probability
of this explanation.
In considering the flower,
the spiral torsion of the ovary,
which brings the lahellnm into
the anterior position, must first
be recognised. The label him
is
tripartite and the larger
middle segment is bifid at its
five end. At the base
of the
labellum a spur is formed by
the bulging out of this seg-
ment of the perianth this ;

serves as the nectary, and the

opening leading into it is situ-
ated just below the gyno-
stemium (Fig. 55:5 A, /?).
The latter bears on the side
that is turned towards tin-

lower lip, and to an insect

alighting on this, the large

FlG. 555. Or'-!
(J nat. si/e.)

\
SECT, ii PHANEROGAMIA 551

stigmatic surface (h) corresponding to two confluent stigmatic lobes. The third stig-
matic lobe is transformed into a structure termed the rostellum (I, k) and stands in
relation to the male organ. The single fertile anther consists of two thecae joined
together by the connective which appears as the end of the gynostemium. The
whole mass of pollen of each of the two pollen-sacs is joined together by an inter-
stitialsubstance which continues below to form a stalk the whole structure, which
;

has a waxy consistence, is called a pollinium, and the stalk goes by the name of
the caudicle. The caudicles terminate below in contact with the rostellum which
forms tough adhesive discs. This relation to the rostellum serves to keep the
pollinia, which lie free in the pollen-sacs, in position, and the adhesive discs attach
the pollinia to any body that comes in contact with them. If an insect alights on
the lower lip and attempts to reach the nectar secreted in the spur, its head or
tongue must touch the rostellum and the pollinia will become attached to it. As
the adhesive discs dry they cause the pollinia to bend forward, so that when the
insect visits a second flower they will be brought in contact with the stigmatic
surfaces.
All Orchids are similarly adapted to insect visitors, though in many the con-
trivances are far more complicated pollination does not take place in the absence
;

of the insects. It should be mentioned that in some forms, e.g. Vanilla, the pollen
remains powdery. Many tropical Orchids are cultivated in greenhouses on account
of the beauty of their flowers.

SUB-CLASS II

14
Dicotylae ( )

The Dicotyledons with few exceptions possess a pair of seed-

leaves these on germination either expand as green assimilating
;

leaves or remain within the seed-ctat and supply the seedling with
the reserve materials stored in their cells. The growing point of the
stem, lying between the cotyledons, grows into the shoot of the
seedling. The main root of the embryo has meanwhile penetrated
into the soil ; as a rule it persists as a tap-root and gives rise to a
regularly branched root-system.
The stem has a circle of open vascular bundles, while the root on
transverse section shows a regularly alternating arrangement of the
xylem- and phloem-groups. The primary meristem situated in the
vascular bundles of the stem, or to the inner side of the phloem in
the root soon becomes completed across the medullary rays and forms
a complete, meristematic ring. By means of this cambium a regular
growth in thickness of the stem and root takes place (cf. Figs. 123-
124, pp. 112, 114; Fig. 128, p. 118; Fig. 138, p. 129; Figs. 146-
154, pp. 138-144).
The typical form of leaf found among Dicotyledons is provided
with a longer or shorter petiole, and often has a pair of stipules
developed from the leaf-base ; a leaf-sheath is usually absent. The
lamina may be simple or compound the latter condition is always
;

2N 1
552 BOTANY PART II

the result of branching during the development of the leaf. The

margin of the leaf presents considerable variety. The venation is as
a rule reticulate (Fig. 556).
The flowers of Dicotyledons exhibit a wide range in their appear-
ance and construction. Most frequently they can be derived from a
type with five whorls of five members ; increase and decrease of the
number both of the whorls and the members are met with. The
Dicotyledons are divided into the two series of the Choripetalae (with
free perianth segments) and the Sym-
petalae (with the petals coherent).

The classification adopted here corre-

sponds in the main to that in ENGLER and
PSABTL's Nat&rliehen Pflanzen-familiei>,\>ul
the grouping together of a number of families
under the older ordinal names has been
retained.
A number of questions raised by HALLIEK
in his recent work on the natural (phylo-
genetic) system of classification of flowering
plants deserve consideration. For instance
the separation of Choripetalae and Sym-
petalac, which HALLIER abandons, is doubt-
less artificial and not altogether satisfactory.
The recognition of the catkin-bearing forms
as reduced and the derivation of the Dico-
tyledons from forms like the Magnoliaceae
has much in its favour. The derivation
of the Monocotyledons from the Polycarpicae
is supported on
<piite different grounds by
E. SARGAKT. While systematic Botany
must endeavour to attain new points of view
Fi<;. 656. Leaf of CVwtalipiM with reticulate,
venation.
and never to regard the prevailing classi-
(} nat. size.)
fication as a sort of noli me tangere,
further developmental studies are necessary before a thorough revision of the
present system can be contemplated. An investigation by STRASBURGER into the
ovule and the formation of the endosperm in Drimys Winteri, a plant which in
the structure of its wood recalls the Gymnosperms, has yielded no indication that
the gap existing between Gymnosperms and Angiosperms can be bridged by the
study of existing forms.

15
Series I. Choripetalae ( )

Order 1. Piperinae
The single Family of the Piperaceae contains a few tropical genera.
Flowers as a rule unisexual and without perianth, associated in spikes ;

typically trimerous but usually reduced. Ovary unilocular, ovule

solitary, basal and atropous. Fruit drupe -like. The embryo is
SECT. II PHANEROGAMIA 553

embedded in a small endosperm

surrounded by a well-developed
perisperm.

The genus Piper includes a large

number of shrubby plants of primaeval
forests.Piper nigrum, from which the
Peppers are derived, is the most im-
portant representative. This is a
root-climber, native to the Malayan
region, but now cultivated through-
out the tropics (Fig. 558). The unripe
provide black pepper, while
fruits
white pepper is obtained from the ripe
fruits after removal of the outer layers
557. Piper cubeba. a, Infructeseence a
;
of the Apericarp. The genus Peperomia
l>,

male flower c, a female flower in longitudinal

;

section d, fruit in longitudinal section. OFFI-

has hermaphrodite flowers.
;

CIAL. (After BERG and SCHMIDT.) OFFICIAL. Piper nigrum and

Fio. 558. I'i[icr nlgnim. (J nat. size.)

554 BOTANY

Piper cubeba (Fig. 557). The latter is a native of Java and is distinguished by the
stalk-like base of the fruit from that of the Black Pepper. It provides CUBKBAK
KRUCTUS.

Order 2. Juglandiflorae

Aromatic woody plants, with anemophilous, unisexual flowers in

catkinate spikes, sometimes containing very few flowers. Ovary bi-

FHJ. 559. Jwglans regia. 1, Branch with young leaves, male catkins and at the tip female (lowers.
male flower,
..',
.i, Female flower. It, Fruit with the outer layer of the periearp in part
removed. (\ nat size.)

carpellary and unilocular with a single, basal, erect, atropous ovule.

Fruit, a drupe. Seed without endosperm.
15li
Family 1. Juglandaceae ( ).- Conspicuous, monoecious trees of
the northern hemisphere with imparipinnate, aromatic leaves arranged
 PHANEROGAMIA 555

alternately. Stipules wanting. Male catkins springing from the

leaf-axils of twigs of the preceding year ; female flowers in terminal

spikes containing more or less numerous flowers; Ovary inferior.

The Wahmt, Juglans regia (Fig. 559), is the best-known representative of the
family. It is endemic in Western Asia and the eastern portion of the Mediter-
ranean region, but the tree is in cultivation throughout Europe. In spring the
axillary buds of the previous season produce long, thick, pendulous catkins bearing
numerous flowers. Each of the latter has 3 to 5 perianth segments, and these together
with the two bracteoles are adherent to the bract and surround the numerous
stamens, which face towards the tip of the inflorescence. The female flowers in
smaller numbers are borne at the summit of the young shoots. The two carpels
terminate in large, feathery, diverging stigmas. The perigone is adherent to the
bract and bracteoles and reaches to the summit
of the inferior ovary. The single loculus encloses
an atropous, basal ovule. Fruit, a drupe. The
exocarp contains abundant tannin. The hard
endocarp is divided into two valves in the plane
of the dorsal sutures of the coherent carpels, the
limits of which are indicated by the partial septum
at the lower part of the fruit. Within the stone
is the embryo, enclosed in a thin seed-coat. The
large cotyledons, which contain oil, are lobed
in correspondence with the false septa that pro-

ject from the inner surface of the ovary. Endo-

sperm wanting. Other species of Juglans and
Carya yield edible seeds and valuable timbers.
Family 2. Myricaceae. Myricn gale is a
dioecious shrub with entire leaves found on moors.
Some American forms yield wax derived from the
fruits (p. 101).
The family of the Casuarinaceae, the syste-
matic position of which is doubtful, may be
mentioned here. They are trees growing by the
sea-shore or on mountains, and are distributed
from southern Asia and Australia throughout
the Polynesian islands. Chalazogamy ( u ) was
first discovered by Treub in Casuai-iim.

Order 3. Salieiflorae

Family Salicaceae. Trees and shrubs

with simple, alternate, stipulate leaves.
Flowers in dioacious catkins, usually
developed before the leaves. Both male
H
and female flowers are naked and stand FIG. 560. Zaliz viminaltf. A, Flower-
ing male twig (nat. si/e) IS, male ;

in the axils of bracts. More or less de- flower with sutitcndiiij; l>i"ict (mo-

veloped scale-like development of the disc llified); i', female inflorescence;

female tlmvers (ma-nilied) F,
or floral receptacle. Ovary of two carpels,
!>!:, ;

fruit (nat. size) G, the same mag-

;

unilocular. Fruit, a capsule containing nified ; //, seed (magnified).

556 BOTANY PART II

numerous, parietal seeds. Seeds without endosperm ;

seed-coat with
a tuft of hairs.

This family is mainly represented in the north temperate zone. Salix, Willow,
and Populus, Poplar, are the only genera. Salix has erect catkins and is adapted to
pollination by insects in relation to this, nectar is secreted by small scales at the
;

base of the flower. Male flowers scented, pollen sticky. The number of stamens
varies from 2 to 5 in the different species. Bracts entire (Fig. 560). Willows occur
commonly by the banks of streams where their shortly stalked, narrowly lanceolate
leaves give them a characteristic appearance. Some species are among the more

Km. 501. I'upulus nigra. 1, Male inflorescence. S, Female inflorescence. ./'. .Male (lower.

A,Female flower. ~>, Fruit. 6', Seed. (1, 3, f nat. size ;.;->;. enlarged.)

abundant plants of high northern latitudes they have subterranean, creeping ;

stems, only the young shoots projecting from the soil (cf. Salix polaris, Fig. 198).
Populus has anemophilous flowers disc cup-shaped no secretion of nectar. The
; ;

long-stalked circular leaves of the Poplars give them a different habit from the
Willows. Flowers similar to those of Salix but with divided bracts. Catkins
pendulous (Fig. 561).
OFFICIAL. SALICIX is obtained from the bark of species of Salix and Populus.

Order 4. Quereiflorae

Trees or shrubs usually with entire leaves and deciduous stipules.

Monoecious. Male flowers in catkins. Ovary inferior ; ovules pendu-
lous. Fruit, a one-seeded nut. Endosperm wanting. Anemophilous.
 PHAXEROGAMIA 557

This order was formerly united with the Juglandiflorae and

Flo. 562. Altiits glutinosa. 1, Flowering branch bearing the small, erect, female catkins and the

pendulous, male catkins. 2, A bract-scale with male flowers. 3, Female catkins. U Female
flower. ;",
Catkin in fruiting condition. 6, Fruit. (1 and the leaf, f nat. aixe ; 2-6, enlarged.)

Saliciflorae in the Amentiflorae ; they include most of our important

forest-trees.

Family 1. Betulaceae. Male flowers adherent to

the bracts ;
in catkins. Female flowers in catkin*.
Ovary bilocular, with two long stigmas a single,
;

pendulous ovule in each loculus.

MOST IMPORTANT GENERA. Alnus glutinosa, the
Alder, is a prominent tree of damp woods, and is also
distributed in swamps and by the banks of streams
(Fig. 562). The leaves are bluntly obovate. The
inflorescences are already evident in the autumn as
stalked catkins, the male long and pendulous, the
female erect and short. Male flowers P 4, A 4 ; a
dichasium of three flowers adherent to each bract
The female flowers are in pairs, their FIG. 563. Alnus glutinosa. Dia-
(Fig. 563).
bracteoles adhering to the bract to form the five-lobed, grams of Fig. 562, 2 and k. Bract
6 ; bracteoles a j3, a' ft', a, /3,.

persistent, woody scale of the cone. Alnus incana

(After EICHLER.)
is distinguished by leaves being grey and hairy
its
below. Betula verrucosa (Fig. 564), the Birch, has a white bark and long-stalked,
668 BOTANY PART II

triangular leaves. "When young, all the parts are covered with numerous glandular
hairs which give the plant an aromatic, resinous odour. The male inflorescences are
formed in the autumn of the previous year, singly or a few together, at the tip of
shoots of unlimited growth. Flowers P2, A 2 in dichasiaof three, adherent to the
;

bract. Anthers deeply bifid (Fig. 564, 3, 4 Fig. 565). Female inflorescences solitary,
;

at the apex of small, short shoots of the current year. Flowers in diehasia of three in
relation to each three-lobed scale the latter is composed of the bract and the two
;

adherent bracteoles. Fruits borne on pendulous catkins winged. After the

;

fruits are shed the scales

of the catkin separate.
Carpinus Betulus, the
Hornbeam (Fig. 566), is
an important forest-tree.
The inflorescences appeal-
in spring, the male, from

axillary buds of the pre-

vious year, either want
leaves or are accompanied
by one or two, the female
are usually terminal. The
bract of the male catkin
bears 4-10 stamens, bifid

Km. oi>4.lielnla vernicosa. 1, Branch with terminal male catkins l-'n.. .v..i. /;<

:uul female catkins on small lateral branches. /. Fi-male flower. Diagrams t'rnin Fig. 564 3
3, Male flower, k, Stamen, 5, A
catkin in fruit. 6, Fruit. and !. b, bract ; a/3, bract-
(1 and 5, g nat. size ; 2-1, and 6, enlarged.) fiiles. (After ElCHLER.)

to the base, but without bracteoles or perigoue. Two female flowers in relation
to each bract ;
each flower with its special bract and pair of bracteoles. The
three latter unite to form a three-lobed involucre which serves as an aid to
distribution by the wind. Corylus avellana, the Hazel (Fig. 567), develops its
inflorescences in the preceding year. The male catkins are freely exposed during
the winter, while the female remain enclosed by the bud-scales, and only protrude
their long red stigmas between the scales at the actual time of flowering. The
male flower has no perianth but has a pair of bracteoles which are adherent
to the bract, as are the four, deeply bifid stamens (Fig. 568). In the short,
female catkins a two-flowered dichasium is present in the axil of each bract

as in Carpinus the fringed involucre also is derived from the coherent brac-
;

teoles and special bract of each flower (Fig. 568). Corylus tubulosa from southern
Europe.
 PHAXEROGAMIA 559

Family 2. Cupuliferae. Inflorescences in the leaf axils, as a

rule of shoots of the current year. Flowers with perianth. Ovary
trilocular, with three stigmas ; two, pendulous, anatropous ovules in
each loculus. One or more female flowers are enclosed in a CUPULE
formed of coherent bracteoles, which only reaches full development

Kic. .")i;i>.
- i
'i'i>in .< tetulus. Branch with male catkins projecting from the buds of the preced-
1,

ing year and female catkins on the growth of the current year. 2, Female catkin in fruit.
3, Male flower, it, Stamen. 5, Bract with two female flowers. G, Female flower. 7, Fruit.

(7, ,.', 7, 3 nat. size ; .!-(! enlarged.)

around the fruit. Another interpretation of the cupule is that it

corresponds to an outgrowth of the axis bearing numerous reduced
leaves (Fig. 569).

MORE IMPORTANT SPECIES. Fayus sylvatiea, the Beech (Fig. 571), is one of our
most important deciduous trees. The leaf is entire, elliptical, shortly stalked,
and, especially when young, covered with fine hairs. Leaves two -ranked.
Inflorescences on shoots of the current season. Male inflorescences lateral, capitate
and pendulous flower with an oblique, bell-shaped perianth and usually 8-12
;
560 BOTANY

stamens. Female inflorescences terminal, capitate and erect ;

flowers in tw<>-

ii.. .V.7. i
..////> iii'tlli' in'. 1, ISniuch \vitli male and female catkins. Group of fruits. J, Bract
.',

with two female Mowers. /t , Male flower. ", Stamen. C, Fruit removed from the cupule.

(7, 2, ft, J nat. size : -l-~> enlarged.)

flowered dichasia. The cupule surrounds both flowers (Fig. 570 B), and completely

L. .'ills.
1'iirylnsavellami. Diagram
of Fi^. 507, a and k. {>, liraet.

o/S, a ft', a, (3,, bracteolcs. (Aftei Flii. &Ci>. Cujnile of (jnei-1-vs AfflHai*. ']>. cuptila :
;//,

EICHLK.I:.) fruit. (Alter DUCHARTHE.)

SECT. II PHANEROGAMIC 561

ft

'
.. 1
"'7 lia^i-anis of tin' female dicha.sia of
1. I :
A, Castnnea vidgaris ; B, Fagus sylvatica ; C, diagram
of the single-flower of Q/vx peihuu'idutu. l>, Uract ; a /S, bracteoles ; o, ft, a ft', bracteoles

of the secondary flowers adherent to the cupnle. (After EICHLER.)

KK;. iXI.Fogii.-i xi/li-<it!i-n. (3 nat. si/c.) J, Branch with male and female inflorescences. -2, Male
flower. .;, Fi-niiilf flower. 4, Open cu])iile witli two fruits. .7, Fruit. 6, Transverse section of
a fruit showing the folded cotyledons of the embryo. (.', .;, n, enlarged.)
2
562 HOT A X Y

envelops the triangular, nut -like fruits;

at maturity it opens by splitting into four
valves. The surface is covered with
numerous, blunt prickles.
Castaiica vulgar is, the edible Che>tnut.
is a native of the Mediterranean region.

The inflorescences on shoots of the current

year bear in some cases only male tlcnuT-..
in others female flowers at the base ami
male flowers above. Flowers grouped in
dichasia. Female dichasia three-flowered
(Fig. 570 A), so that three nuts comr to
be enclosed within the spiny cupule,
Fie.
longitudinal
w hich splits into four valves. The Oaks,
Becttonof the female flower. '/, The young
pedunculata (Figs. 572, 573)
cupuli- f, ovule
; d, ovary
; ; c, perigone ;

/, styl.-: ,,, stixina. (After BERG and

and Qucrcus sessiliftora, are the largest
SCHMIDT, magnified.) deciduous trees of European woods. Leaves

FK.. :,7:i. Quercus yeilniti-nlntn. .1, Klowci-inn l)ranch a male flowi-i (inu-Milii'il)
; /.', C, ; stainrii.-*

(magnified) ;D, a female flower (magnified) /.', iufructescence F, cupule G-Il, seed.
: ; ;
 PHANEROGAMIA 563

oval, margins sinuately lobed ;

those of Q. scssiliftora with longer stalks. The
pendulous, male inflorescences spring at the time that the new foliage is expanding
from axillary buds of the shoot of the preceding year or from the lowest buds
of the shoot of the current year flowers solitary, consisting of a perianth of
;

5-7 segments and 6-12 short stamens. Female inflorescences erect, few-flowered,
in the axils of the upper leaves of the shoot of the current year. Flowers
solitary ; in Q. pedunculata with long stalks, in Q. sessiliftora sessile. Each
flower is invested by a cupule (Fig. 570 0), which is at first inconspicuous, but is
fully developed on the ripe fruit.
The Beech yields firewood, tar, and pyroligneous acid the Oak provides a
;

valuable timber, a bark containing tannin used in tanning, and cork from the
Cork-oak.
OFFICIAL. The GALLS produced on the young twigs of Quercus infectoria as
a result of puncture by the Gall-wasp Cynips tinctoria .Tannic Acid is obtained
;

from these.

Order 5. Urtieinae

Herbaceous or woody plants with small, inconspicuous flowers

closely aggregated in the inflorescence. Mainly anemophilous.
Perianth simple, sepaloid. Stamens equal in number to the leaves
of the perigone and superposed on .the latter. Ovary superior,
composed of one or two carpels, usually unilocular, and containing
a single ovule. Fruit, a nut or drupe. Seeds usually containing
endosperm.

Family 1. Ulmaceae. Trees or shrubs witli simple, asymmetrical leaves

borne in two rows. The pinnate venation, the hairy surface, the serrate margin
of the leaf and the caducous stipules are characteristic. Flowers in clusters in the
axils of leaves of the previous year, hermaphrodite, or unisexual by suppression of
the male or female organs. Perigone of 4-6 leaves, with a corresponding number
of stamens situated opposite to them. Stamens straight in the bud. Unilocular
ovary composed of two carpels with one pendulous ovule.
Ulmus campestris (Fig. 574), the Elm, is a common European tree. The
arrangement of the leaves on the sides of the twigs in two rows and the corre-
sponding branching leads to the leaf-surface exposed by each lateral branch
making a definite angle with the main branch and composing the regular convex
crown of foliage exhibited by older examples. The tree flowers in February and
the fruits ripen before the leaves expand. The fruits are broadly winged and
adapted to be carried by the wind. U. montana, U. effusa are closely related
forms. Several species of Oeltis, in which the fruit is a drupe, are in cultivation.

10
Family 2. Moraeeae ( ). The majority are trees or shrubs
with abundant latex. Leaves alternate, stipules caducous. Flowers
unisexual in globular or disc-shaped inflorescences usually P4, A4, ;

and P4, G(2). Stamens opposite the leaves of the perigone. Ovary
bicarpellary, unilocular, with one, pendulous, anatropous ovule.

IMPORTANT REPRESENTATIVES. In addition to the Mulberry trees, of which

Morus alba is cultivated for the rearing of Silk- worms and J/. niyra as a fruit-tree,
2 1
564 BOTANY

the genus Ficus deserves special mention. The species occurring farthest north
is the Common Fig (Ficus carica, Fig. 575), which is endemic to the Mediterranean
region, and has been long cultivated. It is a low tree with palmately incised
leaves and stipules, which form a cap-like protection to the bud. The inflorescences
are hollow, pitcher-shaped structures with a narrow opening. The flowers are

Km. ~>74. Uhnits ntmpestris (5 nat. size). /, Branch with MI>WCI>. .'. Branch with fruits.

^Sin^le flower, enlarged.?:

borne closely crowded together on the inner surface. The flat, disc-shaped in-
florescences of Dorstenm which bear the flowers on the upper surface, are in many
17
respects corresponding structures ( ). The pollination of the Fig is effected by a
species of wasp, which lays its eggs in the short-styled female flowers or gall-
flowers. The sweet, fleshy portion of the edible Fig is developed from the hollowed
axis of the inflorescence together with the perigones of the individual flowers.
The small, hard, seed-like bodies are the fruits developed from the ovaries of the
small flowers. Some species of Ficus are among the largest trees of tropical forests.
 PHANEROGAMIA 565

The most remarkable is the Banyan

(Ficws benyalensis), which occurs
iu the East Indies. The seeds,
carried by fruit-eating birds, ger-
minate on the branches of trees,
where the plant develops as an
epiphyte. The proper form of the
tree is only seen, however, after
the roots have reached the soil,
and it no longer dependent on
is

the scanty food supply obtainable

in the epiphytic position. The
host-plant is gradually strangled,
additional roots are sent down to
the soil and thicken into pillar-
like supports, and ultimately a
small wood capable of sheltering
an entire villageis developed from FIG. 575. Ficus carica. A, Longitudinal section of
the single small seedling. The an inflorescence. B, Fertile flower. <', Gall-tlower.

D, Male flower. (B-D, enlarged D, after KEENER

; ;
latex of Ficus elastica is obtained
B, C, after SOLMS-LAUBACH.)
from the tree by making incisions
in the bark, and serves as one source of india-rubber. Castilloa clastica is another

Flo. J>76. Hiniiiihi* Instill'*. I, Mali' inflorescence. ._', Female intlorrscnirr. .;. Twu t'rmiili-

flowers in tin- axil of a bract. /<, Cone-like Inflorescences in fruit. (.1 nut. size.)

important rubber-tree of Central America. The gigantic inflorescences wlu-n in

202
566 BOTANY PART n

fruit of species of Artocarpus are eaten raw or cooked and form tin llnad- fruit of
the Tropics. The species of Cecropia are American, myrmecophilous trees (cf. p. 235).
OFFICIAL. The fruits of Ficus carica.
Family 3. Cannabinaceae. Annual or perennial herbs without latex, with
palmately veined leaves and free, persistent stipules. Direcious. Male flowers
P 5, A 5 stamens straight in the bud. The male inflorescences are dichasia, the
;

central branch capable of further growth. The female flowers have a slightly
developed, entire perianth. Ovary bicarpellary, with two large feathery stigmas ;

unilocular, with a pendulous, anatropous ovule. Anemophilous. Fruit, nut-like.

Embryo curved.
GENERA. Humulus lupulus, the Hop, is a native of central Europe; it has a
perennial rhizome, which annually produces a crop of twining shoots (Fig. 576).
The stem and opposite leaves bear coarse hairs, and the former bears hooked
prickles which prevent it slipping down the support. The branches of the female
inflorescence are catkin-like, the scales being formed of the pairs of stipules

belonging to bracts of which the laminae are suppressed. The axillary shoot of
the bract is also suppressed, but each stipule has two flowers in its axil each ;

flower is enclosed by its own bract. These bracts project beyond the stipules
when the inflorescence is mature, and give the latter its cone-like appearance.
Upon them are developed the glandular hairs on account of which the Hop is
cultivated.
Cannabis sativa, Indian Hemp, is an annual herb with palmately divided,
hairy leaves, which are opposite below and alternate in the upper portion of
the shoot. The female inflorescence resembles that of the Hop, but the central
shoot, which in that plant is suppressed, grows out in the Hemp to a leafy shoot.
Only a single flower is present in the axil of each bract. The same process is
repeated in the axil of each leaf of the leafy, middle shoot, so that the whole female
inflorescenceis a repeatedly branched structure. The plant is utilised in Europe
for bast fibres, which are from one to several centimetres long ( 17a ).
its The
glandular hairs which cover all parts of the female inflorescence secrete a sticky
resinous substance which is used medicinally. In the East it is used in the
preparation of a narcotic called Haschisch.
OFFICIAL. Cannabis sativa provides CANNABIS IXDICA. Humulus lupulus
provides LI'PULUS and LUPULINUM.
Family 4. Urticaceae. Perennial herbs or less commonly shrubs. Leaves
simple, stipulate. Flowers unisexual by suppression of parts, as a rule bimerous.
P 2 + 2, A 2 + 2. in the bud, and scattering the pollen when
Stamens inflexed
they suddenly straighten. Ovary consisting of a single carpel, unilocular, with a
basal, atropous ovule. Perianth of the female flower adherent. Flowers in
dichasia, or crowded in dorsiventral inflorescences. Anemophilous.
A numberof the Urticaceae are characterised by the possession of stinging hairs
(cf. Fig. 116), e.<j. the common Stinging Nettles, Urtica dioica and U. uretis, and
the dangerous tropical species of Laportca. Some provide important fibres,
especially Boehmeria nivea from which Ramie fibre is obtained, and, of less value,
Urtica cannabina, and our native species of 1',-tlfn.

Order 6. Centrospermae
Herbs, less commonly small woody plants. Flowers with few
exceptions hermaphrodite and tri- or penta-merous. Perianth and
androecium consisting of one or two whorls. Ovary superior,
 PHANEROGAMIA 567

unilocular, with a single basal ovule or with a free central placenta

bearing a number of ovules.
Family 1. Polygonaceae. For the most part perennial herbs,
with hollow stems swollen at the nodes, and alternate, simple leaves.
The membranous stipules of the latter are coherent to form a
sheath or OCHREA protecting the
terminal bud ; when broken
through by the growth of the
stem this remains as a tubular
sheath around the lower part of
the internode (Fig. 577). The
hermaphrodite flower is typically
trimerous, but a larger number
of members sometimes occurs in
the perigone or the androecium.
Ovary superior, formed of three
carpels, unilocular, with a single,
basal, atropous ovule. Fruit in
nearly all cases a triangular nutlet.
GENERA. Rheum, Rhubarb. This is
an East Asiatic genus, with large, radical
leaves and a large, spreading, paniculate
inflorescence (Fig. 578). Leaves simple,
cordate-reniform, with palmate venation,
sometimes more or less lobed. The
flower has a perigone of two similar
whorls, and two whorls of stamens,
the outer whorl being double by chor-
isis ; P3+3, A6 + 3, G (3). Nectar for

visiting insects is secreted by the large

scales of the disc. The triangular ovary
becomes winged as it develops into the
fruit (Fig. 579). Species of Rheum are
cultivated as ornamental plants and as
vegetables. Rumex acetosa, Sorrel, with
sagittate leaves. The structure of the
flowers of the hermaphrodite species of
Rumex is similar to that of Rheum, but
FIG. 577. Leaf of Polygonum amplexlcaule showing
the inner whorl of stamens is wanting.
the ochrea (st). (J nat. size.)
The species of Polyyonum have a perigone
consisting of five, coloured leaves and a varying number of stamens. The triangular
fruits of Fayopyrum esculentum form Buckwheat (Fig. 508).
OFFICIAL. The rhizome of Rheum officinale, Rh. palmatum, and probably other
species yields RADIX RHEI.

Chenopodiaeeae.
Family 2. Perennial or annual herbs, rarely
small woody with alternate leaves.
plants, Flowers typically
pentamerous, with a single whorl in both perigone and androecium ;

P5, A5, G (2-5). Reduced, unisexual flowers are not infrequent.

203
563 BOTANY I'AUT II

The unilocular ovary contains a basal, campylotropous ovule. Fruit,

a nut. Seed with a curved embryo bent around the perisperm.

Fiii. 5TS. like nut "ffivinule, greatly rcMiiri-d. (After BAILLOS.)

Many of the Chenopodiaceae are strand plants or occur on soils containing a large
amount of salt, such as the great Asiatic salt steppes and deserts 18 ). The Spinach (

(fyiiuicia oleracea) is used as a vegetable. The Sugar Beet (Beta vulgaris, var.
 hi'iim <(/#;"/>. A. Klnuvr ; ;;. tin- saint- cut through longitudinally (', gynaeceum
;

\vitli disc. Hheum compact um : D, fruit. (After LCitssEX, magnified.)

KM;, .wo. - />/" rnt'jarig. 1, Upper portion of the root with

Kic. ">S1. Diagrams of the Caryo-
the rosette of leavt-s, tlie hkulrs of which are cut oft'. phyllaceae. A, Viscaria, septa
lirancli of the inHori'scfiici- (J nat. jui'scnt in tin lowrr part of
1

(Greatly reduced.) '.',

3, Flower cut through longitudinally (' ail ' the ovary ; B, SUene, septa
size).
absent. (After EICHLER.)
enlarged.)
569
570 BOTANY

rapa) (Fig. 580) is a plant of great economic importance. It is a biennial plant, and
in the first season forms a thick, swollen root bearing a bud consisting of a number
of thick -stalked, entire, succulent, and often crisped leaves. From this rosette of
leaves there springs in the second season a highly branched panicle, bearing the
inconspicuous greenish flowers. Ovary formed of three carpels. At the end of

Fit;. 582. Agrostemma Githago. Flowering shoot and fruit (J nat. size). Poisonous.

the first season the root contains cane-sugar as a reserve material, which at this
stage is extracted from the plant. By constant selection the percentage of sugar
is raised from 7-8% to an average of 14% ; it may, however, reach 21-26%. The
original form of the Sugar Beet is Beta patula. Cfienopodium and Atnplex are
common weeds near human dwellings.

Family 3. Caryophyllaeeae. Annual or perennial herbs, very

SECT. II PHANEROGAMIA 571

rarely shrubs, with simple, linear, usually opposite leaves ;

flowers
typicallypentamerous, with calyx and corolla. Two whorls of
stamens, obdiplostemonous. Unilocular or incompletely septate
ovary. K5, C5, A 5 + 5, Gr(5). Fruit, a capsule. Seeds numerous,
embryo curved around the
floury perisperm. In re-
duced forms the fruit is

nut-like (Figs. 581-583). LTO^MMfc-ff

Su H-FA M 1 1,1 KS AND G KNEKA.
1. Alsineae
Calyx poly-
sepalous ; petals with short
claws fruit, a capsule. Gerasti-
;

uni, Sj)er(jula,\\ith pentamerous

flowers ;
Stellaria&ndArenaria
with tri-carpellary ovary. 2.
Paronychieae. Calyx polysepal-
ous ;
corolla wanting or in-
conspicuous ;
fruit nut - like.

Scleranthus, Herniaria. 3.

Sileneae. Calyx gamosepalous;

petals with long claws, fruit
a capsule. Lychnis and A<jr<>-
stemma-witli five carpels, Silene
with three, Dianthus and Sa-
ponaria with two. Cerastium
and Siellaria have white.flowers
with bifid petals and are con-
spicuous, early -flowering forms.
Species of Dianthus, Pinks,
have frequently attractive
colours and scent, and occur in
dry sunny situations. Ayro-
./ niina Githayo (Fig. 582),
Corn-cockle, a hairy plant
is

with pink flowers it is a

;

common weed in corn-fields.

Since its seeds are poisonous,
their mixture with the grain
may have serious results.

Suponariit officinalis occurs KH;. 583.- Sn ,,!,!,-;,!

,ijfi,-;n<ii;.-i($ nat. si/r).
imperially by the banks of POISOJTQUB.
streams it is a herb
attaining
;

the height of a metre, with opposite, broad leaves and rose-coloured flowers, The
saponin contained in all parts of the plant renders it poisonous (Fig. 583).

Order 7. Polyearpieae
This large order includes a number of families of
very different
general appearance but exhibiting as common characters a superior
572 BOTANY

gynaeceum with the carpels free from one another, and a spiral
arrangement of the members of the flower. The tendency to con-
siderable increase in the number of stamens stands in relation
to this.

It however, to be noted that by no means all the plants of the t'uinilirs united
is,

in this order exhibit theabove characters in the same derive. The characteristic
features may indeed be completely wanting, though the existence of intermediate
forms leaves no doubt that the genera in question must be classed here.

Fi<;. 5S4. A'i/ij)/ie vlbu (i nat. size). The spiral arrangement of tin- staim-iis ami |-tals
is shown by their insertions on the ovary to the left.

19
Family 1. Nymphaeaceae ( ). Aquatic plants with submerged or llo.it im;
leaves, the latter often of very large size ; when both types of leaf are present
there is more or less marked heterophylly. Flowers hermaphrodite, with calyx
and corolla. The whorls consist of three (rarely), four, or more numerous
members. Carpels sometimes free. Parts of the flower often spirally arranged
(Figs. 584-586). Latex.
Nymphaea White Water Lily (Fig. 584), has large floating leaves and
alba, the
white flowers, protected by firm green sepals. Within the corolla conies the zone
of numerous stamens and the inferior
ovary composed of numerous, coherent
carpels. The spiral arrangement of the members of the perianth and undmeritim
is seen by the scars of their insertion when they are removed from the inferior
ovary (Fig. 584), and in the floral diagram (Fig. 585). In Nufihar the ovary is
SECT. 1 PHANEROGAMIA 573

superior and the .small petals bear nectaries ; the conspicuous calyx renders the
flower attractive. In the American genus Cabomba the flowers are trimerous and
the pistil is apocarpous. The finely divided, submerged leaves differ in appearance
from the entire, floating leaves. The
carpels are also free in Nehtmbium, both the
leaves and flowers of which are raised above the surface of the water. Victoria

FIG. 585. Nymphaea. !'!.,. :,sii. -.1, Floral diagram. IS, Fruit of (Jubmnlm aquotii-n
Floral (liaf^ram. showing two carpels developed as partial fruits. (At't>-i
HAIULON. x 4.)

reyia from the Amazon, and Euryale ferox from tropical Asia, have gigantic
floating leaves they are often cultivated in Botanic Gardens.
;

The family of the Ceratophyllaceae with the single genus Ceratophyllum may
be placed here. The plants which often occur in masses are submerged aquatics.

Family 2. Ranunculaceae (-). Herbs or rarely woody plants

with alternate, exstipulate leaves. Flowers hermaphrodite, the
members in many cases arranged spirally. Perianth either forming

Fi<;. 08". Flora', diagrams of Ranuiu'iilaceaf. .(, Anemone nemarosa. B, Ailnnix nut unman.*.
i'. Ai-nnitiiiii niijn'llii*. 1> A</iiilrijiu <-/;/n/-ix. l-:,:<"iini<'ifmj ,n<cemai<n. (After EICHLEIS.)

a simple perigone or differentiated into calyx and corolla. Stamens

indefinite. Pollen -grains with at least three places of exit for the
pollen -tubes. Carpels three to indefinite, borne on the convex
receptacle, and forming an apocarpous, superior ovary. Ovules,
borne on the ventral suture, singly or in numbers. The partial
fruits are follicles, achenes, or berries. Seed with a small embryo
enclosed within the large, oily endosperm (Figs. 587-597).
574 BOTANY

IMPORTANT UKXKUA. Many of our commonest meadow and woodland plant*

belong to this order. They are all in greater or less degree poisonous. Numer-
ous species of Ranunculus, characterised by the usually yellow flowers, convex
receptacle, and fruit composed of numerous free achenes, occur in Britain. The
petals have a nectary at the base. Leaves palmately divided more or less deeply.
R. sceleratus is very poisonous (Fig. 590). R. arvensis with large, spiny achenes
or nutlets (Fig. 589). The aquatic species of Ranunculus, belonging to the section

Fi<;. S. Flowci- of Jiunttncitluj! scderatus ; It, tlic sanir. cut tliroujjli longitudinally ;

(After BAILLON.)

Jiatrachium, are often heterophyllous (Fig. 35, p. 34), the floating leaves serving
to support the flowers above the surface of the water.

Species of Anemone are also widely distributed in Europe. A. nemorosa

occurs commonly in woods and is one of our early spring flowers. It has a

horizontal, subterranean rhizome, which terminates in a flower, the further growth

of the plant being carried on by a lateral shoot. Perianth simple, petaloid. All
species of Anemone have at a greater or less distance from the perianth a whorl
of, usually, three leaves forming an involucre (Fig.

591). In A. hepatica this stands just below the

perianth and thus resembles a calyx. All the species
are to some extent poisonous, especially A. Pulsatilla.

Many are valued and are cultivated in quantity

during winter, especially on the Riviera, to supply
the market with cut flowers. The plants of the genus
Clematis are mostly woody and differ from other
Ranunculaceae in having opposite leaves. Many
species are cultivated. C. vitalba is one of our few
native lianes. The achenes of the species of Clematis
and of man y kinds of A mone a provided with hairy
Fii;. .W.i. llanunculus aw ,"(.-.

or leathery appendages, which facilitate their distribu-

Carpel in longitudinal section.
(After HAILLON, enlarge'!.) tion by the wind. The floral axis in the Mouse-tail
is
greatly elongated and the spiral arrange-
(Myosurus)
ment of the small achenes very evident.
upon it is Thalictrum has large, showy
stamens, and the perianth is readily shed (on apogamy in this genus, cf. p. 518).
Calthti palustris, the Marsh Marigold (Fig. 592), is one of the most
conspicu-
ous spring flowers in damp meadows. Perianth simple, bright yellow. Leaves cordate
or reuiform, short-stalked, with erect sheathing base. Fruit composed of follicles.
ffeUeborus foetidus (Fig. 593) has firm, palmately divided leaves. The prominent
inflorescence bears in January or February greenish flowers which stand in the
axils of the broadly sheathing, three-toothed bracts and are spirally constructed
throughout. The five large sepals persist until the fruit is
ripe. The petals,
PHANEROGAMIA 575

(J luit.lsi/.f). Po/sr>fioi;i.
576 BOTANY PART II

KM., .v.il. Anemone 1'ultatilla (J nut. si/f). /'/./ .vo

SECT. II PHANEROGAMIA 577

which are usually thirteen in number, have the form of tubular, two-lipped
nectaries. Numerous stamens. Carpels 3-5, developing into follicles. Helleborus
niger, from the south-eastern Alps, flowers in autumn and winter and is commonly
cultivated. Helleborus viridis. All three species are poisonous. The Monkshood
(Aconitum napellus) is a stately perennial herb with underground tubers and
occurs most commonly in alpine
meadows. The leaves are palmately
divided, the segments being in turn
pinnatelylobed (Fig. 595) and widened
towards the tip. Inflorescence a
dense raceme, reinforced by lateral
inflorescences standing in the axils
of the upper leaves. Flowers zygo-

FIG. 592. Caltha palustris (I nat. sizi-)- FIG. 5'.'3. Helleborus foetidus nat. size).
Potmron. POISONOUS.

morphic. One of the five, dark blue sepals is helmet-shaped, and protects two
long-stalked, tubular, two-lipped nectaries, which correspond to petals. The
remaining petals are wanting or are reduced to inconspicuous, narrow structures
(Fig. 594). Aconitum lycoctonum (Fig. 596) has smaller, yellow flowers of similar
construction. A. Stoerkianum and A. variegatum resemble A. napellus but are
more brightly coloured. All the species are poisonous. Aquilegia, Delphin-
ium and Paeonia are favourite ornamental plants with showy flowers. In Actaea
the fruit is a berry.
2 P
578 BOTANY

OFFICIAL. ACONITI RADIX is obtained from Aconitum napellus. STAI-IIIS-

Fiu. .1114. .1 V.s(nat. size). 1, Flower seen obliquely from in

front. 2,Flower in longitudinal section. 3, The nectaries, formed from
petals, and the androecium after the perigone has been removed. /,, Fruit
composed of three apocarpous carpels. 5, Follicles opened.

Fio. 595. Aconitum napellus (J nat. size.) OFFICIAL and Poisoxovs.

AGRIAE SEMINA from Delphinium staphisayria. CIMICIFUGAE RUIZOMA from

SECT. II PHANEROGAMIA 579

Cinacifuga racemosa. HYDRASTIS RHIZOMA from the. North American Hydrasti

eanadensis (Fig. 597), a perennial herb which sends its subaerial shoots up from
the subterranean rhizome the base of the shoot has keeled scale leaves in two
;

ranks. The and terminate the shoots, each of which bears

flowers are solitary
two foliage leaves. The simple, white perianth falls when the flower opens. The
androecium and the apocarpous gynaeceum consist of numerous members. The
fruit consists of numerous, small berries, each of which includes 1-2 seeds.
The alkaloid HYDUASTINE is obtained from the rhizome.
Family 3. Magnoliaceae. Woody plants from tropical and temperate America
and Asia. The leaves are alternate and usually provided with stipules, which

Fu;. ."I'.Ki. Ai-mtitiDii I

ycoctonum (J nat. size). Po/soxoi'n.

protect the structures in the bud. Flowers and fruit as in the Ranunculaceae,
but the- pollen-grains have only one place of exit for the pollen-tube. Oil-cells in
stem and leaf. Species of Magnolia are largely cultivated on account of their
showy flowers. Liriodendron (Tulip tree) N. America.
OFFICIAL. is obtained from Illicium
ANISI STELLATI FRUCTUS, Star-anise,
anisatum (China). The fruits of Illicium reliyiosum (Japan) are poisonous.
Family 4. Anonaceae. Woody plants of tropical Asia and America. K3,
03 + 3, in other respects like the preceding families. 'Seeds with ruminated
endosperm. Species of Anona yield succulent fruits. A perfume is obtained
from Caiianga odorata.
1
Family 5. Myristicaceae (- ). Tropical, woody plants. Dioecious. Myristica
fragrans (Figs. 598, 599). The male and female flowers are similar in appear-
ance and resemble those of the Lily of the Valley perianth simple, trimerous. ;
580 BOTANY PART II

Male flower with a central column around which 3-15 adherent anthers are
arranged. Female flower with one ovule enclosed by the single carpel. Fruit
berry -like, with coarse flesh, but dehiscent. Seed with ruminated endosperm
and arillus.
OFFICIAL. MYRISTICA, Nutmeg. The seed of Myristica fragrans divested of
its testa.

Family 6. Berberidaceae.
Flowers formed of a varying
number of trimerous whorls.

Carpel {always solitary.

K3 + 3, C3 + 3, A3 + 3, Gl.
In Berberis vulgaris the leaves
on the shoots of unlimited
growth are transformed into
spines ;
the short shoots bear
simple leaves and terminate
in racemes of flowers. The
stamens are irritable to con-
tact (cf. p. 296).
OFFICIAL. PODOPRYLLI
KHIZOMA obtained from Podo-
plujlhim peltatum (Fig. 600),
a North American, herbaceous

plant with large, palmately

divided leaves and a terminal
flower.

Familyl 7. Menisper-
maceae. Similar to the pre-
ceding family but G3. For
the most part tropical, climb-
ing plants.
OFFICIAL. CALUMBAE
RADIX obtained from Jatror-
rhiza palmata (Fig. 601) from
Portuguese East Africa.

Family 8. Lauraeeae.
Aromatic trees or
shrubs with entire
leathery leaves, which
usually persist for several
seasons. Only Sassafras,
FIG. 597. UydrasUs canadensis (J nat. size). The apocarpous!
fruit to the left. OFFICIAL. which has three-lobed
leaves as well as simple
ones, sheds its foliage annually. No stipules. Flowers inconspicuous,
borne in panicle-like inflorescences. They are actinomorphic and com-
posed of trimerous whorls. Androecium of four whorls, some of which
frequently consist of staminodes or are wanting. Anthers opening
by valves. Filaments sometimes bearing glands. Carpels 3, coherent
 m
.. :>W. Mi/ristica fra-
grant, seed, cut through
longitudinally ; g, aril ; h,
outer integument, inter-
Flo. 598. Myristica fragrntis. 1. Twi- \\itli male flowers ( nat. rupted at r by the raphe ;

size). 3,Ripe pendulous fruit opening. 3, Fruit after re- m.ruminated endosperm ;

moval of one-half of the pericarp, showing the dark brown seed m, embryo (nat. size).
surrounded by the ruptured arillus. /,. Kernel treed from the OFFICIAL. (After BERO
seed-coat. OFFICIAL. and SCHMIDT.)

Fio. 600.PodophyIlumpelttittn>, <>FFK:IAI.. (Krom \"t. Pjlanztn/amiUen,

2 P 1
582 BOTANY

to form an unilocular ovary with one, pendulous ovule. Fruit, a

Kir;. i'*\.J<itforrhi:a peltatn. (A nat. size). OFFICIAL. (Att-r UERI; uml .SCHMIDT.)

berry or drupe. Endosperm wanting. Floral formula, P 3 + 3,

A3 + 3 + 3 + 3, G3 (Figs. 602, 603).

IMPORTANT GENKUA AND SPECIES (~).

/. it ii /-us nobilis, the Laurel, is a dioecious, ever-
green tree of the Mediterranean region, which
was well known in the early period of the
Grecian civilisation ;
it is frequently grown
in cool greenhouses (Fig. 604). Large planta-
tions are grown at the Lake of Garda, where
the oil is extracted, and here the trees ripen

their oval, blackish-blue drupes in October.

Kir;. 002. Floral diagram of 1'trgea.
(After EICHLER.) The widening of the floral axis just below the
insertion of tlir- fruit is characteristic. The
gt mis Cinnamomum includes a number of economically important trees such
 PHANEKOGAMIA 583

as theCamphor tree from Japan and China (Fig. 605) and the Cinnamon tree from
China and Ceylon. The latter is a stately evergreen with smooth, leathery leaves
and inconspicuous, greenish flowers in axillary inflorescences. Persea gratissima
(Fig. 602) is a native of tropical Mexico, and is frequently cultivated as a fruit

FIG. 003. Sassafras officinale. After BER<; anil SCHMIDT.) 1, Male inflorescences cm
(J nut. size.
a still leafless Fruits on a leafy shoot. .1, Male flower. It, Female flower. 5, 6,
branch. 2,
Closed stamens of the two outer whorls. 7, Opened stamen of the innermost whorl. H, Ovary
showing the style and the ovule. OFFICIA /..

tree in the tropics. Its fruit is known as the Avocado Pear. Species of Cassytha,
the only genus of the family including herbaceous species, occur throughout the
tropics as parasites resembling Cuscuta.
OFFICIAL. CAMPHORA, Camphor, is 9btained from Cimiamomum Camphora.
CINNAMOMI CORTEX and OLEUM ciNNAMOMi from Cinnamomum zeylanicum.
SASSAFRAS RADIX from Sassafras officinale, a dioecious, North American tree.

2 P 2
FIG. 004 A. Lanfiis nobttis with male
Hovvcrs. (A nat. si/c.)

r,. C04B. Ijaitrns nobilis with fruits.

(J nat. sizi\)

584
 PHANEROGAMIA 585

Fifi. 605. Cinnamomum Camphni'n (., nat. si/c). OFFICIAL.

Order 8. Rhoeadinae

Herbs, or more rarely shrubs, with alternate, exstipulate leaves.

Flowers hermaphrodite, cyclic whorls ;

usually bimerous. Ovary superior, uni-

locular. Placentas on the united margins
of the carpels, projecting more or less
into the cavity. Stigmas commisural, i.e.
situated immediately over the sutures.
Dehiscence of the fruit by separation of
the middle portions of the carpels from
Fl - .-Fi"'- -lia^am . r; to -
the persistent placentas.
*L ., cim. (After EICHLER.)
lamily 1. Papaveraceae. Herbs
with milky juice. Leaves usually pinnately lobed. Flowers actino-
586 BOTANY

morphic. Floral formula, 2, C 2 + K 2, A <x, G(2) or oc. Seeds

with abundant endosperm (Fig. 606)..
The number of the stamens, and usually their position, is indefinite.
In the genera Chelidonium,
Glaucium, Escholtzia, there
are two carpels in Papaver, ;

Aryemont, and J/m//,<///,s-/.sthe

number is greater (14-16).

IMIMM; TANT X \ 1 1\ r. CKNKIIA.

ChelidoniuHi majus, Celandine,
lia> yellow latex and a bicar-

pellarj' ovary. A number of

species of Escholtzia, Argemone,

Fio. 608. Corydalis aureu. u, l':nt

of axis of raceme with a flower ;
b, style and stamens. ( x )

Fid. i>09. Kloial iliii^riiiii of Corydalis

cava. (Affctr EICHI.ER.) At the
base >f tlii- btaMii'ii stainiiii^
i niiiivi'

Fie;. WfJ.Papaver Khoeas (k nat. si/c). <>Ffici.\i.. the spur is a nectary.

and Papaver are cultivated as ornamental plants. Papaver Rhoeas, the Poppy
(Fig. 607), is a common weed in corn-fields or dry meadows. The leaves are
narrow and pinnately divided. The whole plant is covered with coarse, bristly
 PHANEROGAM A I 587

hairs. The bent position of the flower bud is characteristic of many Papaver-
aceae. Ultimately the flower becomes erect as it opens the two sepals forming
;

the calyx are thrown on", and the four, large, wrinkled petals, forming the
corolla, expand. Papaver somnifcrum, which is of oriental origin, has abundant,
white latex. The plant has a glaucous bloom and, except on the flower-stalks,
which bear a few bristly hairs, is glabrous. Leaves sessile, margin irregularly
serrate or lobed. Petals violet or white with a dark patch at the base. Ovary
unilocular, incompletely septate by the projection inwards of the numerous
placentas. Fruit ripens erect on the peduncle. In Papaver the separation of the
central portion of eaeh carpel from the placentas at dehisceuce is limited to the
tips of the carpels. These bend outwards just below the flat stigmatic expansion,
and the kidney-shaped seeds are thrown out of the small openings when the capsule
borne on its long stalk is moved by the wind.
OFFICIAL. Papaver somnifcrum, the Opium Poppy, yields PAPAVERIS CAPSULAE
and OPIUM. Papaver llhoeas yields KHOEADOS PETALA.
Family 2. Fumariaceae. Glabrous herbs, without milky juice, with bipinnate
or tripinnate leaves. Flowers bimerous throughout ;
the number of whorls in the
corolla is doubled. Floral formula, K2, C2 + 2, A2, G(2). The outer petals, or one
of them, bear spurs. In the latter case the flower is transversely zygomorphic
(Figs. 608, 609). The inner whorl of stamens is suppressed. The stamens of the
outer whorl are each tripartite, consisting of a central anther with two thee and
twu lateral anthers each with a single theca borne on a common filament. In
Hir/H'i-niim. the lateral branches join in pairs, and apparently form the inner
staminal whorl, the position of which they occupy. The transversely zygomorphic
flowers of the Fumariaceae with only one spur afford the only example of this type
of symmetry. The fruits of Fumaria are nutlets, those of Corydalis and Dicentra
an- capsules. Dicentra spcctabilis, which is frequently cultivated, has a bisym-
metrical corolla with two spurs. Seeds with endosperm.

3
Family 3. Cruciferae (- ). Annual, biennial, or perennial herbs
without milky juice. Inflorescence racemose, usually without bracts
or bracteoles. Flowers actinomorphic, always lateral, composed of
bimerous whorls. Floral formula, K 2 + 2, C 2 + 2, A2 + 4, G (2).
The outer whorl stands in the median plane ; the four-
of sepals
petals alternate with the sepals. The two outer stamens are shorter
than the four inner ones which stand in the median plane. The
latter correspond to two stamens branched to the base. The carpels
form a superior, usiially pod-like, ovary, which is divided into two
chambers by a false septum stretching between the parietal placentas.
The fruit opens by the separation from below upwards of the main
portion of each carpel, leaving the seeds attached by their stalks to
the central portion formed by the placentas together with the false
septum. Rarely the fruit is indehiscent. Embryo curved. Endo-
sperm wanting or reduced to a single layer of cells coherent with the
seed coat (Figs. 610-616).

The form of the fruit and the position of the embryo in the seed are used in
the subdivision of this extremely uniform family. This general classification, which
dates from the time of Linnaeus, will be used here, but reference must be made to
the more recent, natural division of the family, founded on characters afforded by
588 BOTANY

l-'i... iilo. ('incitera.-. Floral

diagram (/>v,.

Kni.iill. I'linli'iin Klovvri \vitli

perianth removed. (After BAILLON, x 4.)

1)

FIG. 01 Cruciferous fruits. A, Chriranthus cltriri : li, Lepidium sativum ; '', C'upnella bu rua pastoris ;
D, Lunaria birnnis, showing the septum after the carpels have fallen away. K, Crambe mari-
tlma. (After BATLI.ON.)

J77Y7D

Fio. C14. Seeds of Cruciferae cut ucm>s ID

Fin. 613. Tran-iM'i-si- section of the sei-d ot show the radicle and cotyledons. A,
Sratsiea nigra. rail, radicli- <-nt, ciityledons
: ; Cheirnnthiif cheiri ( x 8) B, Sisiim'irtum
:

proc, vascular bundles. (After MOLI.KR.) alliaria(x~). (After BAILLON.)

 PHANEROGAMIA 539

a number of organs, which is given in the Naturlichen Pflanzenfamilien by PRANTL.

LINNAEUS distinguished the Siliquosae, in which the fruit is a siliqua, from the
Siliculosae, in which it is a short silicula. The Siliquosae are further divided into
Siliquosae dehiscentes, with the usual type of siliqua, and Siliquosae lomentaceae, in

I'n;. *'pl5. Cocklearia oJKcinalis, FIG. 616. Brassica nigra (J nat. size).

(After BAILLON.) OFFICIAL.

which the siliqua breaks transversely into one-seeded joints. The Siliculosae are
also divided into Siliculosae dehiscentes and Siliculosae micamentaceae with in-
dehiscent fruits. The Siliculosae dehiscentes were later divided by A. P. DE
CANDOLLE into the S. latiseptae with a bread septum and the S. angustiseptae in
which the septum is narrow.
The number of species and their abundance make the Cruciferae one of our
most important, native families of flowering plants. Their brightly coloured, mostly
590 BOTANY

yellow flowers render them conspicuous in various situations and at all periods of
the year. The nectaries, which are borne on the receptacle at the base of tin-
stamens, also show that the flowers are entomoplulou.s. The family includes a
number of economic plants.
Siliquosae dehiscentes : Cheiranthus Cheiri, the Wallflower (Figs. 612 A,
1.

614 A). Matthiola, the Stock. Numerous species of Brassica have been long in cul-
tivation B. oleracca, the Wild Cabbage, in its various forms
:
(a) sylvcstris, which
occurs on the coasts of Northern Europe and is to be regarded as tin- wild form ; (b)

acephala, Borecole or Kale ;

(c) gonglyodcs, Turnip-

rooted Cabbage (d) gem in i-;

fera, Brussels Sprouts; ()

bauda, Savoys; (f)capitata,
the Cabbage (</) Boti-ijtix,
;

Cauliflower and Broccoli.

Brassica, campesMs, witli
the cultivated forms (a)
annua, (b) oleifera, (c} //</'-

fera. Brassica napus, tlie

Turnip (a) annua, (b) olei-

fera, (c) napibrassica.
Brassica nigra, Black Mus
tard (Figs. 613-616), an
annual plant derived from
the eastern Mediterranean
region, was cultivated even
i n ancient times. The radical
leaves are long-stalked and
lyratc with rounded termi-
nal lobes on ascending
;

the copiously branched

stem they become lanceolate
and gradually smaller. The
plant is glabrous except for
.some bristly hairs on the

upper surface of the leaf.

Inflorescence a raceme ;
tin-

bright yellow flowers stand

out from the main axis, while
KM;. r,i7. Cupixiris spinosa. Flowering branch and a young
fruit borne on the gynophore. the developing fruits an-
(J nat. si/c.)
erect and applied to the
axis. The mature fruits have a short beak the slightly convex valves are keeled
;

and traversed by a well-marked midrib. Seeds spherical. ,S'//i//^/s nibn, White

Mustard, is a hairy .plant, distinguishable from the Black Mustard by the long
broadly-beaked fruits, the valves of which bear coarse bristly hairs. The fruits
project from the axis of the inflorescence. The seeds are yellowish-white and twice
as large as those of Brassica nigra.

Siliquosae lomentaceae
2. Crambe (Fig. 612 E), with the lower portion of the
:

and Cakile are thick-leaved, strand plants. J'tijtliaiiits set tens, the
siliqua sterile,
Radish.
3. Siliculosae dehiscentes latiscptae :
Vesicaria, Aubrietia, Draba, Lunaria,
 PHANEROGAMIA 591

(Fig. 612 D). Cochlearin officinalis, Scurvy Grass (Fig. 615), is a glabrous, biennial
herb which germinates in summer, forming a large rosette of stalked, cordate leaves,
persists over winter, and early in the succeeding summer produces a low, copiously
branched inflorescence of white flowers. The rdsette of radical leaves has dis-
appeared by the time the fruit is ripe.

4. Siliculosae dehiscentcs angustiscptac :

Thlaspi, Iberis with somewhat zygo-
morphic flowers, Capsella (Fig. 612 C).
5. Siliculosae nucamentaceae : Isatis tinctoria, Woad.
OFFICIAL. SINAPIS AI.BAE SEMINA, from Sinapis alba SINAPIS NIGRAE s KM IN A, ;

from Brassica nigra. AHMOUACIAE RADIX, from Cochlearia Armoracia.

Family 4. Capparidaceae. Herbs or shrubs with simple or palmately-divided
leaves, often stipulate. Flowers as in the Cruciferae, K4, C4, A2 + 2, G(2) tliey ;

differ by the gynophore raising the ovary high above the level of
intercalation of a
the other parts of the flower, and by the branching of the stamens (or some of
them) giving rise to an indefinite number (Fig. 617). Fruit usually a berry, in
other cases a capsule or drupe. Seeds without endosperm.
Capparis spinosa is a small shrub occurring on rocky ground in the Medi-
terranean region. The leaves are simple with short, recurved, spiny stipules. The
actinomorphic flowers are axillary and solitary the androecium by chorisis consists
;

of numerous members. The fruit is a berry which reaches the size of a plum and
contains numerous seeds. Capers are prepared from the young flower buds.
Family 5. Resedaceae. Herbs with alternate leaves and zygomorphic flowers
forming a dense raceme. The sexual organs are borne on a gynophore or disc.
K5-8, C5-8, A oo, G (2-6). The^ ovary is unilocular and open above, or the
carpels are free. Reseda odorata, Mignonette. JR. lutea, R. luteola, British.

24
Order 9. Insectivorae ( )

This small order includes the three families of the Sarraceniaceae. Nepenthaceae,
and Droseraceae, which were formerly placed in the Cistiflorse. The name has
reference to the capacity for capturing and digesting insects which is shown by

many representatives of these families (cf. p. 236, Figs. 208-210). The common
affinity of the three families is now generally admitted, and they may perhaps
be connected on to the Resedaceae.
Genera of Sarraceniaceae Sarracenia, Darlingtonia, North American, marsh-
:

and moor-plants.
Genus of the Nepenthaceae Nepenthes, dioecious, climbing, pitcher -plants of
:

tropical Asia (Fig. 46).

Genera of the Droseraceae Dionaea, Venus' Fly - trap, North America.
:

Aldroranda vcsiculosa, a European Water-plant, also found in India and Australia.

Drosera, the Sundew many species occur on peat-moors. Drosophyllum lusi-
;

tanicum, Spain and Portugal.

Order 10. Saxifraginae

Herbs or woody plants of very diverse habit, with actino-
morphic, pentamerous flowers. Androecium usually obdiplostemonous.
Carpels 2-5, free or coherent. Flowers hypogynous, perigynous or
epigynous.
The family of the Podosteinonaceae ( 24a ) may be included in this order. The
vegetative and reproductive organs of these plants are peculiarly modified in
592 BOTANY

relation to their mode of life, and the situations they occupy in the most rapid
currents and waterfalls of rivers and streams in the tropics.
w
Family 1. Crassulaceae. Succulent ( ) herbs or under-shrubs with cymose in-
florescences. Flowers hermaphrodite, with calyx and corolla consisting of a variable
number of members. Obdiplostemonous or haplostemonous. Carpels free (Fig. 618)'

a ft

FIG. 618. Sedum TelepMum. a, Flower ; b, flower in longitudinal section. ( x 4.)

flower hypogynous or perigynous. Scale-like nectaries at the base of the carpels.

Fruit a capsule containing numerous seeds with a small amount of endosperm.
MORE IMPORTANT GENERA. Sedum with pentamerous flowers there are a ;

number of British species. Sempervivum, flowers with from six to an indefinite

number of whorls S. tectorum.
; Bryophyllum with tetramerous flowers, note-

Kit;. 610. Kibet ritbrum. ( nat. size.)

worthy on account of the abundant formation of buds in the indentations ot the

margin of the leaf. Crassula.

26
Family 2. Saxifragaeeae ( ). Herbs or woody plants with
alternate leaves. Flowers hermaphrodite, obdiplostemonous, tetra-
 PHANEROGAMIA 593

merous or pentamerous, perigynous or epigynous. Carpels usually

two ovary uni- or bi-locular. Fruit a capsule or a berry containing
;

an indefinite number of albuminous seeds.

IMPORTANT GENERA. Saxifraga, Saxifrage, small herbaceous plants which are
especially numerous on crags and rocky ground in mountainous districts. They
have a rosette of radical leaves (cf. p. 211, chalk glands) and bear numerous
pentamerous flowers grouped in various types of inflorescence. The two partially
inferior carpels are distinct from one another above. Parnassia palustris is common
on wet moors, K5, C5, A
5 + 5, G(4). Ovary unilocular, placentatiou parietal. One
whorl of stamens modified into palmately divided staminodes, which serve as
nectaries. Cephalotus follicularis is an Australian, insectivorous plant, and possesses,
in addition the ordinary leaves with a flat, elliptical lamina, others meta-
to
morphosed into pitchers. The species of Ribes have an inferior ovary which
develops into a berry, and on this account are commonly cultivated. R. rubrum
(Fig. 619), Red Currant, R. nigrum, Black Currant, R. grossularia, Gooseberry.
Other Saxifragaceae are favourite ornamental plants, e.g. Ribes aureum and
R. sanguineum, Hydrangea, Philadelphus, and Deutzia.
Family 3. Hamamelidaceae. Woody plants of the tropics and sub-tropics.
Liquidambar is mono3cious. Flowers in heads male flowers apetalous, with an
;

indefinite number of stamens ;

female flowers with a perianth and a bilocular ovary.
Liquidambar is a common tree in North American forests.
styraciflua
OFFICIAL. STYBAX PRAEPARATUS from Liquidambar orientalis. HAMAMELIDIS
CORTEX and FOLIA from Hamamelis virginiana.
Family 4. Platanaceae. Stately trees with alternate, three- to five-lobed
leaves and large stipules. Flowers diclinous, monoecious, in small spherical heads,
three or four of which are borne at some distance from one another on pendulous
stalks. Male flowers with three to four stamens
female flowers with four free
;

carpels. Perianth inconspicuous.

Species of Platanus are commonly planted as shade trees by the sides of the
streets.

Order 11. Rosiflorae

27
Family Rosaceae ( ). This is the only family in the order. Herbs,
shrubs, and trees with alternate, stipulate leaves, showing considerable
differences in form and habit. Flowers nearly always actinomorphic,
with the members in whorls. 5, C 5. K
Stamens usually numerous.
Flowers perigynous or epigynous one to many free carpels borne
;

on the expanded, convex or hollowed floral axis. In the epigynous

flowers only the styles are free. The floral axis in many cases
takes part in the construction of the fruit. Seeds usually without
endosperm.
The apocarpous pistil and numerous stamens are characteristic of the Rosaceae

(Fig. 620). Botli these features are also found in the Ranunculaceae or generally
in the Polycarpicae, but the floral members are there spirally arranged while
in the Rosaceae they are in whorls. The latter are further distinguished by the
peculiar development of the floral axis (Fig. 621). In many cases the increase in
number of members of the androecium and gynaeceum proceeds from an intercalary
zone of the hollowed floral axis, and continues for a considerable period. The
2Q
 594 BOTANY

introduction of new members is determined by the spatial relations, so that differ-

ences in the numbers of members arc found in individuals of the same species.
The family is divided into a number of sub-families.

I-',,,, ii-jo. dcnd diagrams of Roaaceae, A,8ortnudowutttM; /', /'/> i'a<ln*; <', /.'K.-K 7,,,,

D, Smtgnism-lia officinalis : /,', Spiraea 7i.v/'< a'-<./"'<<'. (After EICHI.KR.)

() A 5 + 5 or more, Go; other numbers of members in the

Spiraeeae. K5, C5,
whorl are Carpels free or united, each containing two or more
less frequent.
ovules. Fruits usually dehiscent. Floral axis expanded as a more or less Hat disc.
The genus Spiraea has inflorescences bearing numerous flowers many species are ;

FIG. 621. Rosaeeae. Three flowers cut through loiiKitutlinally to show ditt'en-nt forms of
the receptacle. 1, PotentUlu iMlnxti-is ; '. .llrheiitillii uli'ina: '>.
/'//r* muliis. (After I-'OCKI:

in Xttturl. l'fan:enfamilien.)

cultivated as ornamental shrubs (Fig. 623 E). QuiUajn Saponaria (Fig. 622), from
Chili, is an evergreen tree with shortly stalked, alternate, leathery leaves and
terminal dichasia usually consisting of three flowers. The flower has a five-toothed,
nectar-secreting disc projecting above the large sepals. Five of the stamens stand
at the projecting angles of the disc opposite the sepals : the other five are inserted
SECT. II PHANEROGAMIC 595

of the disc. Petals narrow, white. Ovary

opposite the petals at the inner margin
superior. Only tin- middle flower of the dichasium is hermaphrodite and fertile,
the lateral flowers are. male and have a reduced gynaeceum. Fruit star-shaped,
Each carpel dehisces by splitting into two valves.
composed of partial fruits.
Seeds winged.
These are distinguished from the other Rosaceae by their inferior
(6) Pomecu.:
five carpels bound together by the hollow floral
ovary, which usually consists of
receptacle so that only the styles
are free. Each carpel contains one to many
ovules. The fruit resembles a berry, the floral receptacle becoming succulent.
The boundaries of the separate loculi are formed of parchment-like or stony tissue.
The seeds are i.-xalbuminous and enclose a well-developed embryo with fleshy coty-

Kli.. I ,-_'_'. l>nill,ljit >>'/'""((. (* Hilt. Kli.. (>>:'.. /'/// ii.< main*. Flowering slioot,
After A. MKVKK ami SCHCM ANN.) sin-tie flower, ami fruit in longitudinal
section. (J nat. si/c. )

ledons. Apple (Fig. 621, 3 Fig. 623) and P. co'tumunis, the Pear, are
I'lirux Hial-uti, ;

im]M>rtiint and long-cultivated fruit trees, of which numerous varieties are grown.
<
',i,l, ,n in I-II/I/HI-IS, the Quince, has large, solitary, rose-coloured flowers. The fruits
are in shape like an apple or pear, covered with tine woolly hairs and with a pleasant
scent, though not edible when uncooked. In Mespilus yermanica, the Medlar,
the fruit has an apical depression surrounded by the remains of the calyx. The
evergreen Eriobotrya japonica, is commonly planted in the Mediterranean region ;

fruits contain several large seeds, and are succulent, having an acid flavour.
Sorbus (Pyrus) aucuparia, the Rowan. Crataegus (Mespilus) oxycantha, the
Hawthorn, in hedges or planted as an ornamental tree.

(c) Roseae. concave, pitcher-shaped, with one t<> many free

Floral axis

carpels, each of which encloses 1-2 ovules. The partial fruits are nut-like, and are
enclosed by the hollowed floral axis. Rosa with numerous species, varieties and
596 BOTANY

cultivated forms, which have been for long valued 011 account of their form, colour,
and scent (F)g. 620 C). The floral receptacle becomes succulent. Shrubs, some-

Flo. 624. Hagenia al'iistinlcn. 1, Female flower; e, epicalyx : f, calyx :

g, coit)llu(x 4); S, fruit
(nut. .si/e), with enlarged i-picalyx. (After HFKC and Si HMIDT.) Officf.i/..

FIG. 62&. Hagenia abyssiniai. Inflorescence (i nat. size). (After BERC and SCHMIDT.) OFFICIAL.
SECT. II PHANEROGAMIA 597

times straggling climbers, with xmequally pinnate leaves, the large stipules of
which are adherent to the base of the leaf-stalk, and brightly coloured, con-
spicuous flowers. Sepals, inserted on the margin of the receptacle, leafy and
pinnately divided (cf. Fig. 511). The double flowers result from the petaloid
development of many of the stamens. Agrimonia and Hagenia abyssinica have a
dry cup-shaped receptacle. Hagenia is a dioecious tree with unequally pinnate
leaves, the adherent stipules of which render the petiole winged and channelled.
Inflorescence a copiously branched panicle. Each flower has two bracteoles and an
epicalyx. The flowers are unisexual by suppression of the male and female organs
respectively. The corolla later falls off and the sepals become inrolled, while the
epicalyx enlarges. The two free carpels have each a single ovule. Fruit one-seeded
(Figs. 624, 625). Alchemilla has no petals the lack of a conspicuous corolla may
;

lla
perhaps stand in relation to the apogamous development of the embryo ( )

Wi\w
Via. 626.
^PF
Kulin.t ulaeux. Klnwcrin^ branch and fniits. (i uat. size.)

(Fig. 621, 2). Sanguisorba ojficinalis has polygamous flowers (cf. p. 541), without
epicalyx or corolla, aggregated in heads. Flowers tetramerous with 1-2 carpels
(Fig. 620 Z>).

(d) Potentilleac. Floral receptacle plate-shaped, flat or convex and bearing

numerous free carpels which develop into nutlets, drupes, or berries. Each carpel
with one ovule. Epicalyx often present. Potentilla with a number of British
species (Fig. 621, 1). Geum and Dryas have hairy carpels which elongate in fruit
and are distributed by the wind. The latter genus includes arctic and alpine
forms. Fragraria, with small achenes situated on the succulent,
Strawberry,
enlarged, floral receptacle. Ifubus, Blackberry, has numerous species, mostly
scrambling shrubs with recurved prickles. Leaves trifoliate. Ji. idaeus, the
Raspberry, is one of the few species which are not straggling climbers (Fig. 626). The
small drupes are closely crowded on the convex receptacle, only
slightly connected
with one another. The withered style is for a long time visible on the partial fruits.
11. fruticosus.
f9s BOTANY

I'l'/i ni'uc. Flower perigynous, with a single earpfl seated on tha cap-shaped
receptacle (Fig. 620 B). Ovules two. Fruit, a one-seeded <lruje. The group
includes a number of important fruit trees. Pi'inms i-i-i-uatt*. the Wild Cherry
(Fig. 627); P. arivm, (^an : /'. iln,i<>-sti<-n. the 1'hiin : /'. //;/<///<. the A]>ricot,
ind /'.
portion, the I'eaeh, are of Chinese origin :W//s. the Almond, from
/'. ./;/(//</'

the eastern Mediterranean region. The two last-named species have hairy fruits.

Fin. 627. I'ntnvs i-ci'asux (jj nat. si/e). 1, Flowering shoot ttouvr out
; .',
in two (slightly
) : .;, fruits ; /,, fruit cut through loniritudiiuilly.

The succulent mesocarp of the Almond dries up as the fruit ripens, and ruptures,
setting the stony endocarp free.
(/) Chrysobalancae. Frequently with dorsiventral Mowers. A single carpel.
POISONOUS. The seeds of many Rosaceae contain amygdalin, but usually not
in such amount as to be poisonous, owing to the resulting hydrocyanic acid, when
eaten fresh in* small quantity ;
this is, however, often the case with the residuum
the seeds, e.g. of bitter almonds, have been crushed.
left after The leaves of the
Cherry Laurel (Prunus lauroccmsus) may also give rise to toxic etfect>.
OFFICIAL. ROSAF. (t.vi.i.i''.\r. PK IAI.A from cultivated plants ol Jinan i/allicu ;
 PHANKROGAMIA 599

i>LKr>i UOSAK and AC^TA HOSAK from AMYGDALA DULCIS and 7,'o.sv <ln i,ixc, mi,.

AMYCHALA AMAUA from 7V//./1 /.s' <i m

from Primus domesticus.
ii<i<.l.u
I us. PRUNUM
Pnrxi VIKOINIAXAE COUTKX from Prunus serotina. LAUROCERASI FOLIA from
/'/ anus laurocerasus. Cusso from Hagenia abyssinica. QUILLAIAK CORTEX from
Quillaja Saponaria.

28
Order 12. Leguminosae ( )

The common characteristic of all Leguminosae is afforded by the

pistil. This is always formed of a single carpel, the ventral suture of
which is directed to the
dorsal side of the flower
(Figs. 628, 631, 637). It
is unilocular and bears the
ovules in one or two rows
on the ventral suture. The
fruit is usually a pod
(legume), which dehisces by
splitting along both the
ventral and dorsal sutures
(Fig. 642). Nearly all Leg-
uminosae have alternate,
compound, stipulate leaves.
Many are provided with
pulvini (Figs. 236, 237),

Fio. re's. Floral <lia-ram> of Miinos-

acea. .-1, Mimosa pud ten ; I:. io. ii-'. 1
. Acacia senetj'il. Flowering branch (not. si/.').

tiyilmtltllli. (After KlCHLF.B.) (After A. MKYF.R and SCHVMANX).

which effect variation movements of the leaves and leaflets.

Family 1. Mimosaeeae. Trees, and erect or climbing, shrubby

plants with bipinnate leaves. Flowers actinomorphic, peritamerous or
tetramerous. ^Estivation of sepals and petals valvate. Stamens
free, numerous, or equal or double in number to the petals. The
colour of the flower is due to the
length and number of the stamens,
the corolla being as a rule inconspicuous. The pollen grains are often
united in tetrads or in larger numbers. The flowers are grouped in
spikes or heads. Embryo straight in the seed.
600 BOTANY PART II

IMPORTANT GENERA. There are no representatives native to Europe. The

Sensitive Plant (Mimosa pudica) (Fig. 237) occurs as a weed throughout the tropics

Fio. 030. Avn.ela Catechu. ( nat. size. After MEYER auil SCHCMAN.V.)

contact (cf. p. 295). Numerous species of the genus

;uid exhibits great irritability to
Acacia are distributed through the tropics and sub-tropics of the old and new-
worlds some are in cultivation in the Mediterranean region. The Australian
;

forms of the genus are frequently characterised by possessing phyllodes (Figs.

SECT. II PHANEROGAMIA 601

171, 194), the vertical position of which

contributes to the peculiar habit of the
Australian forests. Only the seedlings
of these species have a few bipinnate
leaves which constantly succeed the coty-
ledons. Some American species of Acacia
are inhabitedby ants (cf. p. 235, Fig. 205)
which live in the large stipular thorns
and obtain food from Belt's food-bodies at
the tips of the pinnules. The plants in
this way are protected against the attacks
FIG. (531. Floral diagrams of Caesalpiniaceae.
of leaf- cutting ants. Many species of
A, Cercis sUiquastrum ; B, Tamarindus
Acacia are of considerable economic value
indica. (After EICHLER.)
owing to the presence of gums and tannins

Kic.. r.:i'A Cassia angustifolia. (5 nat. size. After A. MEYER and SCHUMANN.) OFFICIAL.
602 BOTANY

in tin- i-ortex or in the heart-wood. A. >;,/,',/,,/ (Fig. 630) ami -/ .-/'/ an- Ka-t
Indian trees from which Catechu is obtained.
OFFICIAL. By the disorganisation of the parem-hynia of the stein of A<->-<
xi-nfijiil (Soudan and Senegamhia) and of other species, AI-ACIAK <;TMMI
is ohtainrd.

Tliis exudes from wounds as a thick fluid and hardens in the ,-iir.

Family 2. Caesalpiniaeeae. Trees or shrubs with pinnate or

bipinnate leaves. Flowers usually somewhat dorsiventral. Corolla

(* nat. si/<>. Al'ti-i A. MKYKK and SCHI MAXX.)

-
OFF/CIA/..

with ascending imbricate aestivation (Fig. 631) or wanting. Typical

floral formula: K
5, C 5, A 5 + 5, Gl. The number of petals and
stamens is often incomplete. Embryo straight.
IMPORTANT GENERA. examples of the Caesalpiniaeeae are
Characteristic
afforded by the flowers of Cassia. In the sub-group Seniw, to which C. a/i.<tusti-
folia belongs, the sepals and ]>etals are both five in number and free (Fig. 632).
 SKCT. II 1'H. \\KKOGAMIA 603

Tlie lower overlapping petals aresomewhat larger than the upper ones. Of the
ten stamens the three upper ones are short and sterile while the other seven, the
filaments of which are curved and convex below, diminish in length from above
downwards. The anthers open by means of terminal pores. The pod is com-
pressed and broad and Hat. The (lowers are borne in racemes in the axils of the

KKI. ii34. TaiiKiritiiliis in-

dica. Fruit in longitud-
inal section. M, the fleshy
mesocarp. (Aft<>r BER<; <.. i>3"'. -i'n/,/i'/j','i'ii I.Hiiiixilnrffii. (J Mat. size) witli a single flower
and SCHMIDT.) Orfn-i.u.. (nat. size) and tlie fruit (reduced).

leaves of the shrub, which is about a metre high. The bright green, equally
pinnate leaves have small stipules at the base (Fig. Tamarindus indicn 632).
(Fig. 633) is a handsome tree, native to tropical Africa, but now planted throughout
the tropics. Its broadly spreading crown of light foliage makes it a favourite
shade-tree. The racemes of flowers are terminal on lateral twigs bearing equally
pinnate leaves. The individual flowers are markedly zygomorphic (Fig. 631 B}.
The bract and two bracteoles are soon shed. The four sepals have a somewhat
 604 BOTANY PART II

FIG. 637. Floral diagrams of Papilionaceae.

A, Vtcia Faba ; B, Cytisvs Laburnum. (After EICIILER.)

FIG. 636. Krameria trinndra. (f nat.

size. After A. MKYER andi SCHU-
MANN.) OFFICIAL.*

Fit;, das. i.<4ii* eornleutadu (4 nat.

si/i'X Flowering shoot flower, keel,
;

stamens. Carpel (nat. size) and fruit Fi<;. {W. Julan Pereirae. (g nat. size. After BERG
(J nat. size).
and SCHMIDT.) OFFICIAL.
SECT. II PHANEROGAMIA 605

cruciform arrangement. The three, red-veined, yellowish-white petals stand erect.

The three stamens are coherent at the base and lie close to the easily movable
curved carpel, so that the anthers and stigma project together. The i'ruit is
peculiar. The pericarp is differentiated into an outer brittle exocarp, a succulent
mesocarp, and a firm endocarp consisting of stone-cells investing the more or less
numerous seeds individually (Fig. 634). Humboldtia laurifolia has hollow inter-
nodes inhabited by ants (Fig. 206, p. 236). The almost imperceptibly dorsiventral
flowers of Copalfera (Fig. 635) have no corolla ;
the four sepals are succeeded by
8-10 free stamens. The fruit is one-
seeded but opens when ripe. The seed
is invested on one side
by a succulent,
irregularly limited arillus. None of the
Caesalpiniaceae are British. Cercis sili-
quastrum from the Mediterranean region,
which bears its flowers on the old woody
stems (cauliflorous) (Fig. 631 A), and
Gleditschia triacanthos (N. Am.) are
sometimes cultivated as ornamental
plants in gardens.
OFFICIAL. SENNA INDICA, the pinnae
of Cassia angustifolia (Trop. East Africa

FIG. 640. Myroxylon Pereirae. See IVxt. (En- l-'n;. (141. Fruit of Myraxylvn Pereirae
larged. AfterBKRO and SCHMIDT.) OFFICIAL. (3 nat. size). OFFICIAL.

and Arabia, cultivated at Tinnevelly in Southern India) SENNA ALKXANDRIKA ;

from C. acutifolia ;
Cassia fistula (Trop. Am.) yields CASSIAE PULPA COPAIBA is ;

obtained from Copalfera Langsdorfii and other species TAMARINDUS from the ;

succulent mesocarp of Tamarindus indica HAEMATOXYLI LIGNUM, the heart-wood

;

of Haematoxylon campechianum (Trop. Am.); KKAMERIAE RADIX from Krameria

triandra, a shrub growing in the Cordilleras. Flowers atypical ; the sepals
brightly coloured within the corolla small.
;
Three stamens opening by pores at
the summit. Fruit spherical, prickly. Leaves simple, silvery white (Fig. 636).

Family 3. Papilionaeeae. Herbs, shrubs, or trees with, as a rule,

imparipinnate leaves. Flowers always markedly zygomorphic.
Calyx of Corolla of five petals, papilionaceous, with
five sepals.
descending imbricate aestivation (Fig. 637). Stamens 10; filaments
606 KOTANY

either all coherent into a tube surrounding the pistil or the posterior
stamen is free. Seeds with a curved embryo.

component parts of a papilionaceous flower arc seen separately in Fig. 638.

Tlu-
The which overlaps the others in the hud, is termed the standard
posterior petal,
(vexillum). The two adjoining, lateral petals are the wings (al;e!. and the two
lowest petals, usually coherent liy their lower margins, together form the keel

Ki<;. <<-l-. i.'iitixus Isoljurnum (J nat. .size).

(carina). The upper ends of the stamens are usually free and curve upwards, as
does also the style bearing the stigma.
iMi'oiiTAXT SUB-FAMILIES AND GENERA. The (1) Sophoreae and (2) Podalyrieae
approach the Caesalpinaceae in having the stamens more or less free. The former
is characterised
by pinnate leaves, the latter by simple or palmate leaves. The
genus Myroxylon, belonging to the Sophoreae, is' of importance on account of the
balsam obtained from secies belonging to it. Myroxylon Pcreirac is a tree of
moderate height with alternate, imparipinnate leaves (Fig. 639). The flower- an
 I'HA.XKKOliAMIA 607

borne in terminal racemes and have a large vexillum, the other petals remaining
narrow and inconspicuous. The stamens are only coherent at the base, and bear
conspicuous, reddish -yellow anthers (Fig. 640). The fruit very peculiar. The is

ovary has along stalk and hears two ovules near the tip. One of these develops
into the seed of the indehiscent, compressed pod, which has a broad wing

along the ventral suture and a narrower wing along the dorsal suture (Fig. 641).
The bell-shaped calyx persists
on the stalk. 3. Gcnistcac.
All ten stamens are coherent
(Fig. 637 B).Leaves pinnate
or simple, margin entire. Genista,
Sarotliamnus, Lupin us, Cytisus
(Fig. 642). The Laburnum
( Cytisus Laburnum) is one of the
commonest ornamental trees of
our gardens and grows wild in
the Alps. It has tripinimtf
leaves and long pendulous
racemes of yellow flowers. C.

Adami, which is found occasion-

ally in gardens, was supposed to
have arisen by the grafting of
C. jiurjiurcus on C. Lului-raitm.
The racemes and Mowers of one
and the same individual sonic-
times resemble the former and
sometimes the latter ancestral
form (Fig. 247). The inter-
mediate form is however sterile,
and probably corresponds to an
ordinary hybrid grafted on C.
Laburnum. Ulcx, Furxc, a char-
acteristic British plant. Spar-
> in
in, distributed in the Mediter-

ranean region. 4. Trifolieae.

Leaves trifoliate, serrate.

Flowers aggregated in heads or
in spikes or umbels. Stamens
(9) + 1. Indehiscent fruits.

TrifiiiittiH, Clover, with persist-

ent calyx and corolla. Metlicago,
Medick, with deciduous corolla ;
Klii. -7 I-IIIIK! iis Att>-i
li-}.'!.; . I
//iiniiiiijii. (1 tint. sixe.
fruit sickle-shaped or spirally \. M \ 1:11 ami Si in OFFICIAL.
i
MANN.)
twisted. Mililotus, Melilot,
with racemose inflorescences. Trigonclla. Ononis, Rest-Harrow. The increase
in the amount of nitrogen in the soil effected
by the root-tubercles (cf. p. 232,
Fig. 204) of Legumiuosae iinds its practical application in European agriculture
in the cultivation of species of TrifoUu'm, Afeilicayo, and 5. Lotcac.
Lupinus.
Lotus, Bird's-foot Trefoil Leaves imparipinnate, lowest pair of
(Fig. 638).
leaflets owing to the absence of the petiole
resembling stipules. Anthyllis,
Kidney-Vetch. Tctrayonolobus. 6. Galeyeae or Astragaleae. Leaves impari-
608 BOTANY

pinnate ;
a very large sub-family, the plants belonging to which show much
diversity in habit. In species of Astragalus, which are low shrubs of the eastern

Fio. 6M.Glycyrrhiza glabra (J nat. size). OFFICIAL.

Mediterranean region and of western Asia, the rhachis of the leaf persists as a
sharply pointed thorn for years after the leaflets have fallen. These spines serve to
protect the young shoots, leaves, and flowers (Fig. 643). Our native species are
herbaceous. Robinia is an American tree of rapid growth with very brittle wood,
SECT. II PHANEROGAMIA 609

which is often planted and known as False Acacia. Glycyrrhiza, Liquorice, is a

native of S. Europe (Fig. 644). Colutea, Caragana, Indigo/era. Wistaria sinensis
is a cliraher with beautiful blue flowers, often grown against the walls of houses.

7. Hedysareac. Distinguished by the jointed pods, in which the seeds are

isolated by transverse septa. Coronilla (Fig. 645). Ornitkopus sativus, Bird's-
Foot. Arachis hypogaea, Ground-nut, an important, oil-yielding fruit of the tropics
and sub-tropics.After flowering the flower-stalks penetrate the soil in which the
fruits ripen. 8. Vicieae. Leaves

with terminal tendrils, corre-

sponding to the terminal leaflet ;

the leaves may thus appear to

be paripinnate. The cotyledons
remain within the seed-coat
and do not become green.
Stamens Ficia, Vetch
(9) -f 1. ;

Pis-urn, Pea
Ervum, Lentil
; ;

Lathyrus, Everlasting Pea.

Faba, the Broad -Beau,
is an
erect without tendrils
plant, ;

the terminal leaflet is reduced

to a bristle-shaped
stump. 9.
Phaseoleac. Twining plants
with tripinnate leaves. Coty-
ledons usually come above
ground and turn green. Stamens
(9) +1. Pkaseolus, Kidney
Bean. Physostigma.
POISONOUS. Among our
common Leguminosae only Cyt-
iaus Laburnum is
extremely
poisonous. The other species
of the genus, e.y. C. alpinus, C.
purpureus, 0. Wcldeni, and C.
lifloms are poisonous, but are
more rarely met with than the
ordinary Laburnum. Coronilla FIG. 645. Coronilla varia (nat. PoisOA'OCS.
size).
varia (Fig. 645), with umbels
of rose-coloured flowers, and Wistaria sinensis are also poisonous.
OFFICIAL. Astragalus gummifer and other species yield TRAGACANTHA.
(ii.vt YiiUHiz.YE RADIX is obtained from
Glycyrrhiza glabra. Spartium scoparium
yields SCOPARII CACUMIXA. Physostigma venosum, a West African climber,
yields PHYSOSTIGMATIS .SKMINA, Calabar Bean. Andira araroba, a Brazilian
tree, contains a powdery excretion in cavities of the stem called ARAROBA ;
if KYSAROBINUM is obtained
from this. The heart-wood of Pterocarpus santalinus,
an East Indian tree, is PTEROCARPI LIGNUM. KINO is obtained from the
juice
flowing from incisions in the trunk of Pterocarpus marsupium. Myroxylon
tuluifcra (S. America) yields BALSAMUM TOLUTANUM, and M. Pereirae (San
Salvador) BALSAMUM PERUVIANUM.

2 R
610 BOTANY

Order 13. Gruinales

The flowers of the majority of the plants belonging to this order

are hermaphrodite, pentamerous, and radially symmetrical, with a
superior, septate ovary. 5, C 5, 5 + 5, (5).K When the A G
flowers are zygomorphic they frequently exhibit reduction. Stamens
coherent at the base, obdiplostemonous or haplostemonous. Nectaries
to the outer side of the stamens or as a circular disc within the
stamens. Ovules, usually pendulous ; micropyle directed upwards.
The Terebinthinae are included in this order as above defined. Their dis-
tinguishing characteristic is the intrastanrinal, annular nectary.
Family 1. Geraniaceae. Herbs or small shrubs with, as a rule, alternate,
stalked, palmately veined 'leaves. Flowers
conspicuous, solitary in the leaf axils or form-
ing cymo.se or umbellate inflorescences. They
are usually ^ctinomorphic in Pelargonium ;

zygomorphic and provided with a spur.

Stamens 10 or f>. Two ovules in each loculus.

l'i<.. i'.-ir.. Floral diagrams of Geraniaceae. A, 6>;r<,,M/ KM.. 47.---Kniit of I'llnriiimlKni

prattiise ;B, Pelargonium zonalc. (After EICHLER.) V. (After BAII.I.OS.)

When ripe the five beaked carpels separate from a central column, and either open t>
liberate the seeds, or remain closed and by the hygroscopic movements of the awn-
like portion bury the seed in the soil (Figs. 646, 647. Cf. Fig. 222, p. 271).
Geranium with ten stamens and Erodium with live stamens occur in Britain.
Numerous species of Pelargonium from the Cape are cultivated as ornamental

plants the peculiar scent of the vegetative organs

;
is due to the ethereal oil

secreted by the glandular hairs.

Family 2. Tropaeolaceae. Tropaeofum nmjus, Indian Cress, climbs by help
of the petioles. Flowers zygomorphic, spurred, with 8 stamens and 3 carpels.
Fruit, a capsule. Leaves peltate (cf. Fig. 191).
Family 3. Oxalidaceae. Leaves compound, in sonic species irritable to
mechanical stimuli (liinpltijt.um. Oxalis acetosff/", Wood Sorrel, cf. p. 295).
Family 4. Linaceae. Herbs or shrubs with alternate, narrow, entire leav.-.>.
Flowers actinomorphic. Stamens coherent at the base those opposite the petals ;

developed as staminodes. Ovary 5-locular, each loculus incompletely divided

by a false septum in each chamber is one pendulous ovule. Fruit a capsule
;

(Fig. 648 A-C). Liiiiini utttati$ttmUTn, Flax, has long been in cultivation. It is
 PHANEROGAMIA 611

an annual, and bears numerous blue flowers, which last only a short time, in
racemose cincinni. The flower has five, free styles. The stem bears numerous
small, narrow leaves. The bast-fibres after proper preparation are woven into
linen. The seeds yield oil.
OFFICIAL. LINUM, the seeds of Linum usitatissimum.
Family 5. Balsaminaceae. Succulent herbs, with translucent stems, some-
times swollen at the nodes, and simple leaves. Flowers zygomorphic with five
free stamens. Capsule
dehiscing elastically. Im-
patiens noli tangere, I.

parvifiora, Bal- and /.

samina. Regarding the

occurrence of cleistogam-
ous flowers in Impatiens
cf. p. 518.

Family 6. Erythroxy-
laceae. Erythroxylon
Coca is a small Peruvian
shrub, with entire, simple

Fid. 648. Linum usitatissimum. A,

Flower ; B, androecium and gynae-
ceum ; C, capsule after dehiscence.
(A ,
nat. size
; B, C, x 3). OF-
FICIAL. F;<;. 6W.Erythroxi/lon Coca. nat. size.) OFFICIAL.
(|j

leaves and axillary groups of small, white flowers. Petals with appendages.
Stamens 10, united at the base into a tube, G (3), but only one loculus of the
ovary is fertile. Fruit, a one-seeded drupe (Fig. 649).
OFFICIAL. From E. coca COCAE FOLIA, which yield COCAIXA, are obtained.
Family 7. Zygophyllaceae. Floral formula typically K5, C5, A 5 + 5
G(5);
with an intrastaminal disc.
OFFICIAL. Guiacum
qfficinalc, a "West Indian tree with opposite, paripinnate
leaves. Ovary bicarpellary, bilocular. Fruit winged. It yields GUIACI LIGNUM
and GUIACI RESINA.
Family 8. Rutaceae. For the most part shrubs or trees, rarely herbs, with
2 R 1
612 BOTANY

alternate, pinnate leaves. Flowers usually actinomorphic, tetramerous or penta-

merotis with a large iutrastaminal
disc. Oil-containing cavitii- in

all the members.

I M l'i 1 11 1A NT ( i KN KUA. Rlltd
graveolcns (Fig. 650), the Rue, is

a somewhat shrubby plant with

pinnately divided leaves. The
terminal flowers of the dichasial
inflorescences are pentamerous in
robust examples all the other ;

flowers are tetramerous. Dictam-

nus Fraxinella has panicles of
conspicuous, dorsiventral flowers ;

the carpels are free in their upprr

portions. The important genus
29
Citrus has peculiarly con -
( )

structed flowers (Figs. 651, 652).

The numerous stamens are uiiitnl
in bundles and arranged in a

Fio. 651. Floral diagram of.Ci<) i

vulgarif. (After EKHI.EB.)

nat. size).

single whorl. The number of

carpels is also increased. The
fruit is a berry the succul-
;

ent portion is formed of large

with abundant cell-sap
cells
which project into and fill
up the loculi of the ovary.
The leaves of many species
are simple and provided with
more or less winged petioles.
Other species have trifoliate
leavesand the articulation at
the base of the lamina shows
that the apparently simple
leaves correspond to impaii-
pinnate leaves, of which only
the terminal leaflet is devel-
oped. The thorns at the base
of the leaf are derived by
modification of the first leaves Fio. 652. Citrus vulgaris (A nat. size). OFFICIAL.
SECT. II PHANEROGAMIA 613

of the axillary bud. Citrus is originally an East Asiatic genus ; a number of

species inhabit the warmer valleys of the Himalayas. All the important cul-
tivated forms have been obtained from the Chinese. Citrus decumana, the
Shaddock, is
tropical C. inedica is the form which was
;
known to the Greeks in
the expeditions of Alexander as the Median apple. It is now widely spread and

Fin. 053. QIMSSM amara. (Xat. size. After BERC and SCHMIDT.) OFFICIAL.

has a number of varieties of which Citrus (nied-ica) Limonum is the Lemon.

This tree was introduced into the Mediterranean region in the third or fourth
century. Citrus (medico) Bajoura has thick-skinned fruits from which citron is
obtained. Citrus Aurantium occurs in two distinct forms, C. (Aurantium)
vulgaris and C. (Aurantium) sinense. The latter is a variety of the Bitter
Orange tree of apparently Chinese origin, and already comes more or less true to
seed. The name Orange, which is introduced into many languages and is
derived from the Sanscrit word Nagarunga, points to the ancient cultivation of
 614 BOTANY

the plant in India. The name was first applied to the Bitter-fruited Orange,
which found its way west much earlier that the sweet variety, and was sub-
sequently applied to the latter. Citrus nobilis, the
Mandarin, is also of Chinese origin.
OFFICIAL. Citrus Aurantium, var. Bigaradia, yields
AUKANTII CORTEX SICCATUS, AVRASTII CORTEX RECEXS,
and AQUA AURANTII FLORis. Citrus medica, var. limonum,
gives LIMOXIS CORTEX, and LIMONIS succus. BUCHU FOLIA
are obtained from Barosma betulina. CUSPARIAE CORTEX
from Cusparia febnfuga. JABORANDI FOLIA from PH-
carpus jaborandi, a tree-like shrub with large, imparipinnate
leaves, native of Eastern Brazil.
Family 9. Simambaceae. K5, C5, A 5 + 5, G5.
Carpels free. No
Contain bitter principles.
oil glands.
OFFICIAL. QUASSIAE LIGXVM from Picrasma excelsa
(West Indies) and Quassia amara (Surinam). The latter
(Fig. 653) is a small tree with beautiful leaves and showy
flowers.

Family 10. Burseraceae. Woody plants with resin

passages. Fruit, a drupe. Commiphora abyssinica and C.

FIG. 054. Floral diagram of Polygda myrtifolia. (After EICHI.KR.)

Fio. 6a5.l'olygala Senega. A, Flower ; a. small 'b, large sepals

; c,
;

keel ; e, lateral sepals ; d, androeciuiii. D, androecinm h, .anthers

; FIG. 656. Polygala Stnega.
(magnified). (After BERG and SCHMIDT.) OFFICIAL. (J nat size). OFFICIAL.

Schimperi are trees found in Arabian East Africa. Boswellia Carteri and B. Bhau
Dajianae are small trees (3) from the same region which yield OLIBANUM. Canarium.
OFFICIAL. MYIIRHA, Myrrh, from Balsamodendron Afyrrha and other species.
Family 11. Polygalaceae. Herbs or shrubs with small simple leaves and
SECT. II PHANEROGAMIA 615

dorsiventral flowers. K5, C3, A(8), G(2). The two lateral sepals are petaloid.
Three petals, the lowest of which forms a keel. Stamens 8, coherent into a tube.
Anthers opening by pores. Ovary bilocular, formed of two carpels a single ovule ;

in each loculus (Figs. 654-656). Polyyala chamaebuxus is a small shrubby plant

occurring in the Alps. P. vulgaris and P. amara occur in Britain.
OFFICIAL. Polygala tieneya (North America) yields SENEOAE RADIX.

31
Order 14. Trieoceae ( )

The
large Family Euphorbiaeeae which belongs to this
of the
order is of much greater importance than the small family which is
here placed along with it. The plants belonging to the Euphorbiaeeae
are of very diverse
habit. The order in-
cludes herbs, shrubs,
leafless succulent
plants, trees with nor-
mal foliage, and others
with scale leaves and
assimilating phyllo-

Kic.. 057. Ovule of Euphorbia FIG. 658. Mercurialiti annua (i nat. si/.'). Male plant in
divica showing the obturator. flower and single male flowei-. Portion of a female plant,
(Att'r PAX in Knglrr-J'runtl.) single female flower ami fruit. I'OISOXOUS.

clades. The plants

agree, however, in possessing unisexual, actino-
morphic flowers, with a simple perianth or with no trace of the
latter. Androecium diplostemonous or stamens numerous. The
female flowers are especially characterised by the superior, trilocular
ovary formed of three carpels in each loculus are one or two
;

pendulous ovules with a ventral raphe, and the micropyle directed

upwards arid outwards.
The micropyle covered by a placenta! outgrowth called the obturator (Fig.
is

657) ; and nourishing the pollen-tube, and disappears

this assists in conducting
after fertilisation. The CARUNCULA, which is formed from the outer integument
(Fig. 662 D), persists on the other hand in the seed the separation of the latter
;

from the placenta is assisted by it.

2 R 2
 616 BOTANY

The fruit is a capsule, the outer walls of which contract elastically

away from a central column, and thus open the loculi.
IMPORTANT GENERA. Many Euphorbiaceae are ditecious or IIIOIKH inii.s, and have
flowers of relatively simple construction. Thus Mercurialis (Fig. 658), two species
of which occur in Britain, is characterised by its bicarpellary ovary. Croton is a
tropical genus including valuable official
plants. C. Eleuteria and C. Tiglium. In
the Spurges (Euphorbia), of which there
are several British species, a number of

!'!<;. MO. Diagram Of a ilichasial liranrh nf

Ma, with three cyathia, only the inicldl <.t i.

which has a fertile female flower. (After EICRI.F.I:.)

Fui. 651'. Euphorbia ciiparifniii* O nat. si/e). l-'n.. iir.1. />;./.. i///m ntM/era. (Nat. si/e.
PoffiOXot'x. After BEIU: and SCHMIDT.)

the extremely simply constructed flowers are grouped in a complicated inflorescence

termed a CYATHIUM (Figs. 659-662). This consists of a naked, terminal, female
flower, borne on a lung bent stalk surrounded by a number of groups of male flowers.
Each of the latter is stalked and consists of a single stamen, the limit bct\\e.-n
which and the flower-stalk is distinguishable. In some cases the female flower
and each male flower are provided with a small perianth- The whole cyathium,
which is an inflorescence, is always enclosed by five involucral bracts alternating ;

with these are four nectar-secreting glands, the presence of which increases the
likeness between the cyathium and a flower. The fifth gland is wanting, and the
 PHAXEROGAMIA 617

inverted female flower hangs down in the gap left. Between the groups of male
flowers, which stand opposite to the bracts (Fig. 660), are branched hairs which are
visible when the cyathium is cut through longitudinally (Fig. 662 B). The cyathia
are usually grouped in dichasia, and these in turn form an umbellate inflorescence,
with three to many branches. It often happens that the female flower is only

developed in some of the cyathia, remaining rudimentary in the others. Many

species of Euphorbia, especially the African species, are succulent-stemmed plants
resembling Cacti in general appearance (cf. Fig. 661).
Euphorbia, like many but not all the other plants of the family, contains a
milky juice, which is secreted in non-septate latex-tubes. This juice, which in
many cases is poisonous, exudes wherever the plant is wounded.

KK:. i
id-.'.
l-:nj, !,,,,!,
:
T.iitlnirit. A, Cyathium (x .~0.
t i /;. C.vathium cut thruuf;h longitudinally
(x 7).Fruit alter dehiscence showing the centv;il column (<). /', Seed in longitudinal
C,
section showing the embryo embedded in the eiid<|>enn <", caruncuKi (x 4).
; (A-D after
BAILLON.)

An
important constituent of the latex of species of Hevea (If. Uiebcri, discolor,
lutea, guyancnsis, Spruceana) is CAOUTCHOUC.
rigidifolia, pattoifolia, As Para
Rubber obtained in the tropics of South America, especially in the Amazon region,
this affords about one half of the total rubber supply. In addition Manihot
Glaziovii, another South American plant of this order, which yields Ceara Rubber,
must be mentioned. A nearly related plant, Manihot utilissima, provides in its
tuberous roots a very important food in the tropics. The starch obtained from
these roots forms mandioc or cassava meal, the finest varieties of which, as tapioca
or Brazilian arrowroot, are of commercial importance. The shrub, which is a
native of Brazil, is now cultivated throughout the tropics.
Ricinus communis (Fig. 663) is a tall shrub of tropical Africa. In our climate
it is annually killed by the frost. The hollow stem bears large palmately divided
leaves. The terminal inflorescences (Fig. 664) are overtopped by vegetative lateral
618 BOTANY PART II

branches. The male flowers, situated towards the base, have a membranous calyx
of 4-5 sepals, enclosing the branched stamens the end of each branch bears a;

theca. The female flowers, nearer the summit of the inflorescence, have 3-f> sepals
and a large tripartite ovary. The latter is covered with warty prickles, and bears
three large, bifid, red stigmas. In each loculus of the fruit is a mottled seed with
a whitish caruneula. Mallotut

Kii;. i',63. Ricinux i-'i

-iitly iviluct-d. (After HAILLON.) 1'oisoxoi'S and OFFICIAL.

OFFICIAL. Croton Eleuteria (Bahamas) yields CASCAKILLA. C. tiglium (East

Indies), OLE CM CKOTOMS. OLEUM KK-IXI, Castor Oil, is obtained from Jliciniis
communis.
The Callitrichaceae are a small family of water plants, with naked, unisexual
flowers. Stamen 1, carpel 1.

Order 15. Sapindinae

This order im-hidcs trees and shrubs of very different appearance, with cyclic
flowers. The latter exhibit reduction in the perianth, androecium, and gynaeceum.
SECT. II PHANEROGAMIA 619

One or two ovules are contained in each chamber of the ovary ;

when the ovules
are erect the raphe is ventral, when they are pendulous it is dorsal ;
this differs
from what is the case in the Euphor-
biaceae.

Family 1. Buxaceae. Buxus

sempcrvirens, the Box, is a poisonous,
evergreen shrub with very close
wood, native of Central and Western
Europe. In the axils of small, de-
cussate bracts stand capitate in-
florescences, consisting of a terminal,
trimerous, female flower, and a
number of tetramerous male flowers.
Family 2. Auacardiaceae. Dis-
tributed in the tropics and warm
temperate regions. Ovary of 1-3

carpels. Fruit usually a one-seeded

drupe. Schizogenous resin passages.
Mangifera indica, an important fruit
tree of the tropics. Species of Rhus
yield Japan lacquer. R. toxicoden-
dron, a North American climbing

FIG. 665. Floral diagram otllexaiiiiif"liinii.

(After BICHI.ER.)

FIG. 664. Ricinus cnmmunis. Inflorescence (J

nat. si/e) ; young fruit cut through longitudin- l'i... i.y.t>. Floral diagram of Euonymus
ally. OFFICIAL. europaea. (After EICHLER.)

shrub, is extremely poisonous. Species of Pistacia (Mediterranean region)

have
pinnate leaves. P. vera is cultivated on account of its drupaceous fruits called
Pistachio nuts.
Family 3. Aquifoliaceae. Ilex aquifolium, the Holly, is an evergreen shrub
or tree of Western Europe (Fig. 665). K4, C4, A4, G(4). In each loculus of the
620 BOTANY PART II

ovary is a pendulous ovule. /. paraguariensis in extra-tropical South America

yields Paraguay Tea or Mate.
Family 4. Celastraceae. Euonymus europaea, the Spindle tree (Figs. 666,
667), is a British shrub with inconspicuous flowers; K4, C4, 4, G(4). The A
stamens are inserted at the margin of a large disc two ovules in each loculus.
;

In autumn the bright red arillus of the seeds, which hang by their stalks from the
opened pale red capsule, makes the fruit conspicuous.
Family 5. Aceraceae. Trees with opposite leaves. The flowers show a
tendency to become unisexual by abortion of stamens or carpels. Disc intra- or

Ki<;. 0<5~. Kunnymus europaea. A, Flowering branch (reduced) ; B,

a flower (magnified) ;

C, D, the fruit (nat. size). (B-D, after ESGLER-PRANTL.) Poisosocs.

extra-staminal. Carpels two. Fruit, a winged schizocarp (Fig. 668). In Britain

Acer campcstre and A. pseudoplatanus. The characteristic lobed outline and
palmate nervation of the leaves, as a rule, enable the genus to be readily recog-
nised.

Family 6. Sapindaceae. Tropical trees or shrubs usually with obliquely

zygomorphic flowers with an extra-staminal disc.
The crushed seeds of Paullinia cupana, a liane of tropical Brazil, yield GUARANA.
Family Hippocastanaceae. Closely related to the Sapindaceae. Acsculus,
7.

Horse-chestnut. Acsc. Hippocastanum, the White Horse-chestnut, is a tree of

hilly regions of the eastern Mediterranean (Fig. 485 B). Aesc. pavia, the Red
 PHANEROGAM I A 621

Horse-chestnut, is a native of the mountains of the Atlantic side of North America.

Both are commonly cultivated.

FIG. 66S. Acer psevdoplatanus (J nat. size). 1, Branch with pendulous terminal inflorescence.
2, Male flower. 3. Female flower, k, Fruit. (2 and -^'.enlarged.)

Order 16. Frangulinae

Shrubs or trees with inconspicuous, actinomorphic, tetramerous or
pentamerous flowers. There is only one whorl of stamens, which
are antipetalous. Flowers hypogynous or perigynous. Ovary
composed of two to four carpels with one or two erect ovules in each
loculus. Disc intrastaminal.
622 BOTANY PART II

Family 1. Rhamnaeeae. The only native genus of this family,

which is distributed in the tropics, is Ehamnus, the Buckthorn (Figs.
669-671).

A o B o

KM;. tiTO. Hlif in nun Frangula. Flower

cut through longitudinally, a, Re-
Fir;. 6<5!>. Floral diagrams of \A, Rhamnus cathartica ceptacle ; 6, calyx ; c, petal d, a
;

(represented as hermaphrodite) and B, Rh. stamen e, pistil (magnified). (After

;

Frangula. (After EICHLER.) BERG and SCHMIDT.)

Rh. Frangula, the Berry-bearing Alder, is a shrub with alternate, entire leaves
provided with small stipules. The flowers are solitary or in groups in the axils of
the leaves. K5, C5, A5, G(2). The* floral receptacle forms a cup-shaped disc.

Fin. ffi'l. nti" vinifera. Opening

flower. Calyx b, corolla c,
, ; ;

disc d, stamens e, ovary (mag-

; ;

niiied). (After BERG and SCHMIDT.)

FIG. 671. Rhamnus frangula (i nat size). Flowering FIG. 673. Floral diagram of Ampel-
branch and portion of a branch bearing fruits. opsis hederacea. (After EICHLER.)

Two (less commonly three) carpels stigma undivided. Fruit, a drupe with two
;

or three seeds. The wood was formerly used as a source of charcoal for the manu-
facture of gunpowder. Rh. catharticus has usually spiny branches bearing
SECT. n PHANEROOAMIA 623

Flowers tetramerous throughout, diceciou.-,

opposite leaves with serrate margins.
by suppression of stamens or carpels female flower with four free styles and a
;

four-seeded drupe. Seeds with a dorsal furrow. Colletia spinosa and C. crueiata
are leafless South American shrubs the thorns of the former are cylindrical, those
:

of the latter flattened laterally.

OFFICIAL. Ithuimm.x fiin-fliiunus yields CASCARA .SAURADA or RHAMNI
Ft'RSHIANl COKTEX.

Family 2. Vitaeeae. Shrubby plants climbing by means of

tendrils ;
leaves palmately lobed or divided. Flowers hypogynous.
Ovary composed of 2-4 carpels and with a corresponding number of
loculi. Two ovules in each loculus. Disc often consisting of
separate glands (Figs. 672, 673).
IMPORTANT GENERA. J'itis, climbing shrubs of the northern hemisphere.
Vitis vinifera, the Grape Vine, is a cultivated plant with numerous races and
varieties. The tendrils correspond to shoots and stand opposite to the leaves ;
they are at first terminal, but become displaced to one side by the development of

the axillary shoot. The whole shoot is thus a sympodium. The original relation
between the tendrils and the axillary shoots, which are both recognisable at the
growing point, cannot be completely followed in the ontogeny. The inflorescence
is a panicle taking the place of a tendril ;
intermediate forms between inflorescences
and tendrils are of frequent occurrence. Calyx only represented by a .--mall rim ;

Corolla thrown off when the flower opens. Currants art- the seedless fruits of Vitis
viniferu, var. apyrena. Species of Ampeloptit distributed in North America and
Asia go by the name of "Wild Vines some of them have tendrils with adhesive
;

disc (Fig. 29, p. 27). Cissits is a large, exclusively tropical genu.-.

OFFICIAL. UVAE, Raisins from 1'itis rin(f>','n.

Order 17. Columniferae

The essential character of this order is afforded by the androeciuni

of the regularly pentamerous, actinomorphic, hermaphrodite flowers.
One of the two whorls of stamens, usually the outer one, is sup-
pressed or only represented by staminodes, while the other whorl
has undergone a greater or less increase in the number of its
members by chorisis. The branching is frequently accompanied by
cohesion of the filaments. The carpels also sometimes exhibit an
increase in number as a result of branching. The superior ovary is
then divided into a corresponding number of loculi.
Family 1. Tiliaceae. For the most part trees or shrubs, less
commonly herbs, with simple, stalked leaves provided with deciduous
stipules. Calyx polysepalous. ^Estivation of calyx and corolla
valvate. Stamens completely free from one another with introrse
anthers. A 5 + 5, or only the inner whorl is present and has usually
undergone branching (Fig. 674). Ovary with two to many loculi,
and one to many ovules in each loculus. Style simple.
Most of the genera are tropical. The herbaceous species of Cot-chorus yield
Jute. In Britain two species of Tilio, Lime, occur. These are stately trees with
624 BOTANY PART II

two-ranked, j>etiolate leaves, the stipules of which are soon shed. The leaves,
which have a serrate margin, are asymmetrical. The inflorescence (Fig. 675 A)
is coherent with a bract for half

its length ;
this serves as a wing
in the distribution of the fruit.
A second bract at the base pro-
tects the axillary bud, which will

expand in the succeeding season.

Below tin- terminal flower are
three small hracteoles ;
the low-
est of these is sterile while the
two upper ones have single
r,
flowers in their axils. Each of
these flowers a pair of bracte-
lias

oles which serve as the subtend-

ing leaves for further branching.

The umbel-like inflo'rescence of
the Lime is thus composed of
dichasia ;
TIJin platyphyllot lias

A
Kx;. ii75. Til i ulnii/olia. A, Inflorescence (a), with
bract '<
(nat. size). If, Longitudinal section of
frnit (magnified); u, pericarp; p, atrophied dissepi-
074. Tiliaceae. Floral diagram ment and ovule ; </,
seed r, endosperm ; s, embryo ;
;

(Tilia). (After EICHLER.) t, its radicle. (After BERG and SCHMIDT.)

3-7, T. parvifolia 11 or more flowers in the inflorescence. The hairy ovary has two
ovules in each of its tive loculi. The fruit only contains one seed (Fig. 675 2?).
32
Family 2. Sterculiaceae ( ). Tropical herbs, shrubs, or trees,
with simple leaves and deciduous stipules.
Flowers with a gamosepalous calyx. Corolla
twisted in the bud. Stamens coherent to form
a tube ; the antisepalous stamens are stamin-
odial, the antipetalous stamens are often in-
creased in number. Anthers extrorse.
'^sas^sst^
\ ;. r,76. -Sterculiaceae.
IMPORTANT GENERA. The most important plant is
Floral diagram (Then- _, . -..
-r.- - ,..t~x T^ . ,
the Gocoa tree (Theobroma Cacao, Pigs. 6/6, 6/0- It
bra*,,). <Aft.T EICHLEK.)
is a native of tropical Central and South America, but

has long been cultivated. The increasing demand has led to its being grown with
more or less success in many tropical colonies. It is a low tree with short-stalked,
firm, brittle, simple leaves of large size, oval shape, and dark green colour. The
young leaves are of a bright red colour, and, as in many tropical trees, hang limply
SECT. II PHANEROGAMIA 625

Fio. 6"". Thi'iiin-iiiiiii 1'in-iiii. 2, Stem bearing froita. 2, Flowering'branch. 3, Flower. 4, Circle:of
stamens. .7, Stamens from anterior side. (.;, .',, about nat. size; 5, enlarged; 1, 2, greatly
redneed.) OFFICIAL.

downwards. The on the main stem or the

flowers are borne
older branches, from dormant axillary buds (CAULI-
and arise

FLORY). Each petal is bulged out at the base, narrows con-

siderably above this, and ends in an expanded tip. The form
of the reddish flowers is thus somewhat urn-shaped with five
radiating points. The pentalocular ovary has numerous ovules
in each loculus. As the fruit develops, the soft tissue of the
septa extends between the single seeds the ripe fruit is thus ;

unilocular and many-seeded. The seed-coat is filled by the p, $*$ Malvaceae

embryo, which has t\vo large, folded, brittle cotyledons. Cola Floral diagram <

2s
626 BOTANY

He a ,,/inata and C. vera, natives of tropical Africa, yield the Kola nuts which are
used in medicine.
OFFICIAL. Theobroma Cacao, from which OLEUM THEOBROMATIS is obtained.

Family 3. Malvaceae. Herbs or shrubs, frequently with pal-

mately-lobed, stipulate leaves which in the young condition often have
a velvety covering of stellate hairs. Flowers with an epicalyx, a
gamosepalous calyx, and a corolla which is
usually of considerable
size and attractively coloured, and is contorted in the bud. Pro-
tandrous. Stamens united into a tube around the ovary ; the free
ends of the stamens, each of which bears a single reniform theca,
project from the margin of the staminal tube. Pollen-grains with

Fio. 679. Flower of Althaea officinalis, cut

through longitudinally. a, .Outer fi, ;

inner calyx ; c, petals d, androecium'; /,

;

pistil t, ovule (magnified).

; (After HERO Fio. 680. Malfn tttvtttrit. a, Flower; b, flower-
and SCHMIDT.) bud ; c, fruit (nat. size.)

spiny exine (cf. Fig. 474). Carpels usually numerous. Fruit, a

schizocarp or a capsule (Figs. 678-681).

IMPORTANT SUB-FAMILIES AND GENERA. 1. Malveac. Schizocarps composed

of numerous carpels arranged in a circle. Malva, with number of British sprrirs
;>.

(Fig. 680). Perennial herbs, with long-stalked, palmately-veined leaves. Flowers

solitary or in small cymose inflorescences in the axils of leaves. Three, free
segments of the epicalyx. Petals usually rose-coloured, deeply notched. In
Althaea the whole plant is clothed with stellate hairs giving it a soft velvety
appearance. Epicalyx of 6-9 segments united at the, base.
2. Fruit, a capsule usually formed of five carpels with a correspond-
Hibisceae.
ing number of loculi. Hibiscus is frequently cultivated as an ornamental phmt.
Gossypium, shrubs with three- to five-lobed leaves with long stalks. Flowers with
a large epicalyx of three segments, which completely covers the calyx. Fruit of
three to five carpels, loculicidal. Seed covered with long hairs which aid in its
dispersion by the wind. When stripped from the seeds and cleaned these hairs
 PHANEROGAMIA 627

form cotton wool. The most important species of Cotton are G. barbadense,
G. arbor eurn, G. herbaceum (Fig. 681).
OFFICIAL. Gossypiurn barbadense and other species yield GOSSYPIUM.

l-'n;. I'lM. Flowering branch and open fruit ot Oossypium herlxwcum (J nat. size). OFFICIAL.

Order 18. Cistiflorae

The plants belonging to this order are characterised by their

usually regular, pentamerous flowers ; the stamens are increased in
number by chorisis, or when the separation of the branches is incom-
plete they form distinct bundles ;
the superior ovary is
usually
trimerous.

Family 1. Ternstromiaceae. Herbs, shrubs, or -trees, with alternate, simple,

leathery leaves. Flowers regular ;
aestivation imbricate sometimes with indistinct
;
628 BOTANY TAUT II

separation of calyx and corolla. Stamens numerous. Ovary septate. Thea

chincnsis (Fig. 682) is a small evergreen tree or shrub, native to southern China.
The young leaves and tips of the
shoots yield TEA after being fer-
mented and dried. The tea-plant is

cultivated on an extensive scale in

China, Japan, and many tropical
colonies. CAFFEINE is obtained from
Thea ch inensis. Th ea japon icn th e ,

Camellia, is a favourite ornamental

shrub.
Family 2. Guttiferae. Herbs.
shrubs, or trees with opposite, simple,
entire leaves. Flowers sometimes
diclinous. Stamens in bundles.
SchizogenoOB intercellular spares
containing secretions in all parts of
the plant. In Britain .//////<//</>/;<.
with a number of .-pecies stamens :

in three or five bundles (Fig. 683).

Oil glands apparent as translucent

spots in the leaf -blade. Gardnia

Mangostana, the Mangosteeu, is one
of the most esteemed fruit-trees of
the Asiatic tropics.
OFFICIAL. darcinia Haulm mi
isa tree of Farther India, with thick
leathery leaves and diclinous flowers.
FIG. 682. Thea chinensis. Flowering shoot ( nat. The male flowers have numerous
stamens seated on a short globular
receptacle the female flowers have an ovary composed of three to several carpels
;

surmounted by a broadly peltate stigma. The red[contents of the secretory organs

which exudes on wounding the plant provides when dried, CAMBOGIA, Gamboge.

I'm. OS3. H/II* li'-i'iii

ti'ti-ii]it<-i-iiiit. a. Flower, somewhat '. -Flor;il dia.uiam of Heli-
maxnitit'd ; I,, fruit ; ]>, tin- dried, iii-rsistent petals. until* mum riiltjuiT. (Cistacrai\)
(X 2.) (Atl.-i KICIII.ER.)

Family ''. Dipterocarpaceae. Contain secretory reservoirs. Characterised by

the great enlargement of some or all the sepals after fertilisation. l>riil>nliiiii>?.

i, yields i'.orneo camphor. Dammar ( :K! ) is obtained from Shorm II' i-

SECT. II PHANEROGAMIA 629

FIG. eS5.Helianthemum vulgare (i nat. si/el.

Family 4. Cistaceae.
Perennial herbs or
shrubs with opposite,
simple, stipulate leaves.
Pentamerous, regular
Mowers, with numerous
stamens and three or five
carpels united to form a
unilocular or multilocu-
lar ovary with a single
style. In Britain the
Rock Rose (Helianthe-
mum vulgare) ; the pros-
trate stem bears small
elliptical, sessile leaves
and racemes of bright
yellow flowers (Figs, 684,
685). Many species of
Cistus are characteristic
shrubs of the type of
vegetation of the Medi-
terranean region, known
as the Maqtiis.

Family"). Violaceae.

FIG. 687. Vida tricolor. A, Entire itlant (ivduced); st, stipules.

/;. a stamen, enlarged; c, connective: nt, nectary. C, gynae-
FIG. r>80. Floral diagram ceum, enlarged ; n, stigma. D, transverse..section of ovary.
of Viola. K, fruit (nat. size).
630 BOTANY PART II

Herbs, .shrubs or trees, usually with dorsi ventral flowers with only five stamens.
Ovary unilocular with a simple style. There are a number of species of Viola in
Britain. The flowers have the anterior petal prolonged backwards as a spur, into
which two neetar-secreting processes of the two anterior stamens project (Figs. 686,
687). Many kinds of Violet have, in addition to these conspicuous flowers adapted
to insect pollination, inconspicuous, cleistogamous flowers which are self-fertilised
r
(cf. p. 309). ioia tricolor, the Wild Pansy, has large pinnately-divided stipules
J

hardly smaller than the leaf-blade itself.

Order 19. Passiflorinae

Flowers radial, perigynous or epigynous. Ovary of three carols, unilocular,

with parietal placentation usually three bifid stigmas.
;

Family Paasifloraceae. Plants

climbing by means of tendrils.
Flowers large, complicated by out-
growths from the floral axis (corona
and disc). Passiflora, Passion
Flowers ornamental plants from
;

South America.
Family Caricaceae. (,'"//<"

papaya, the Pa paw, has in its

latex a ferment (pa]>ayotin) re-

seinbling pejisin. Tropical.

Family Begoniaceae. Herl
with obliquely cordate leaves.
Flowers unisexual. Ovary inferior.

Mostly tropical. Many spei-ie> of

Begonia are in cultivation.

34
Order 20. Opuntinae ( )

Family Cactaceae. For the

most part leafless plants with suc-
culent stems, natives of America. In
size they range from very small to

gigantic forms. Flowers hermaphro-

dite, actinomorphic, less commonly
dorsiventral. Perianth of mam
Fio. 688. Cereusgemnetrizans. Two of the rib* or
member8j sl ra u y arranged and
,i

ridges of a five-ribbed stem bearing flowers and fruits

(| nat. size). showing a gradual transition from
the calyx to the corolla. 8tamen>
and numerous.
carpels Ovarv inferior, unilocular, with numerous parietal
placentas. Ovules with long stalks. Fruit, a berry, the succulent tissue being
largely derived from the stalks of the seeds.
Peireskia and some species of Opuntia possess leaves. Other species of Ojntiitin
have flattened branches. Cereits, Echinocactus, with longitudinal ridges on the
stem Mamillaria has free projections (mamillre). The numerous groups of spines
;

on the shoots, ribs, or separate mamillae correspond to axillary shoots, the sub-
tending leaves of which are reduced while the leaves of the expanded axis of the
axillary shoot are metamorphosed into spines (Fig. 688).
SECT. II PHANEROGAMIA 631

Cactaceae form a dominant constituent of the vegetation in the dry south-

western regions of the United States and in Mexico. They are also widely dis-
tributed in the West Indies and South America. A similar habit is found in some
Euphorbiaceae and Asclepiadaceae living under corresponding climatic conditions.
There are numerous epiphytic Cactaceae, especially species of Rhipsalis, Epiphyllum,
and Phyllocadus, which clothe the branches of trees and affect the general
aspect of the vegetation.
Opuntia ficus indica has become naturalised in the Mediterranean region. The
fruits of this species and of
others of the genus are edible,
and the plants are cultivated
as fruit-trees.Some Cactaceae,
such as Anhalonium, contain
highly poisonous alkaloids and
saponine. The Cochineal insect
isgrown upon species of Opuntia
and Nopalea (N. coccinellifera).

Order 21. Thymelaeinae

Shrubs or trees, with simple,

entire leaves, often closely
crowded. Flowers perigynous,
hermaphrodite, actinomorphic,
tetramerous or pentamerous.
Corolla often wanting. Stamens
in one or two whorls. Carpel,
one. Ovary with a single ovule.
Family 1. Thymelaeaceae.
Ovule pendulous. Daphne
Mezereum (Fig. 689) isa poisonous
shrub, possibly native to Britain,
which flowers in February and
March before the leaves appear.
The flowers are rose-coloured,
scented, tetramerous, and have
no corolla. The leaves form a
close tuft until the axis elongates.
The fruit is a bright red berry.
FIG. 68!i. Daphne Mezereum (J nat. size). OFFICIAL
and Poisuxiti'ti.
In the Alps and in the Medi-
terranean region there arc numerous species of Daphne, all of which are poisonous.
OFFICIAL. Daplnn' Mi^:reum, D. Laureola, and D. Gnidium yield MEZEKEI
CORTEX.
Family 2. Elaeagnaceae. Ovule erect. Hippophae. Elaeaynus. The leaves
and young twigs are covered with shining peltate hairs.

Order 22. Myrtiflorae

Leaves opposite, simple, entire, exstipulate; flowers hermaphrodite,
actinomorphic, tetramerous or pentamerous, hypogynous, or at most
632 BOTANY PART II

perigynous. Ovary septate. Increase in number of the stamens is

frequent.

Family 1. Lythraceae. Lythrum salicaria. Purple Loosestrife. K6, C6, A6

+ 6, G(2). Heterostyled with three forms of flower (ef. p. 311).
Family 2. Melastomaceae. An exclusively tropical family, easily recognised l>y
the beautifully shaped and regularly veined leaves. Favourite hothouse plants.
Family 3. Onagraceae (
:!3
). K4, C4, A 4 + 4, G(4). Androecium obdiplo-
stemonous. Ovary inferior (Figs. 690, 691).
MOKE IMPORTANT GENKIIA. Epilobium, Willow-herb, with numerous spcch * :

the fruit a capsule, and the seeds have hairs serving for wind- dispersal.
is

Ocnothera. The power of mutating possessed by plants of this genus was recognised
by DE TRIES and forms the experimental basis of his hypothesis of mutation.
Circaea, Enchanter's Nightshade.
Trapa, Water Nut. Many forms
are cultivation, for instance
in
the species of Fuchsia, in which
the calyx is petaloid. These
plants are natives of America.
Fruit, a berry.
Family 4. Halorrhagi -

daceae ( 35a ). Aquatic plants

with small, inconspicuous flowers.
Myriophyllum. Hippuris.

Km. ti90. Floral diagram of Oe>inth> TH

(Onagraceiie). Kit;. 031. Fiichxic Flowers (nat. si/e).

36
Family;".. Rhizophoraceae ( ). Plants occurring in the .Mangrove formation
along tropical coasts, characterised by vivipary and the possession of stilt-routs m
i-es]iiratory roots. These adaptations are related to the peculiarities of the
situations in which the trees grow,
llhizophora (Fig. 692) Bruguicra Ceriopr. ; ;

Family 6. Combretaceae. TcrminaUa Chebula yields HTBOBALAN8.

Family Evergreen shrubs or trees; leaves opposite,

7. Myrtaeeae.
leathery, often Flowers actinomorphic, tetramerous or
aromatic.
pentamerous. Androecium of many stamens, which are often arranged
in bundles which have
originated by branching. Carpels two to
many, united with the floral axis to form the inferior ovary. Fruit,
usually a berry or a capsule.

IMPORTANT GENERA. The Myrtle (Afyrtus communis), which occurs in the

Mediterranean region, is the only European species. Species of Eucalyptus, from
SECT. II PHANEROGAM I A 633

Australia, especially E. ylobulus, are commonly planted in Italy, on account of their

rapid growth and useful timber. Young plants have opposite, sessile leaves, but
older trees bear stalked, sickle-shaped leaves which hang vertically. The shadeless
condition of the Australian forests formed by these trees, depends in part on

Ki',. i'i'. i-2.

l!hi;iiplinr<' ODHJvgata (J nat. size).

this character, but is partly due to the distance apart of the individual trees.
E. a unjrjdalina,which reaches a height of 155 m. and a circumference of 30 m. at
tin base of the trunk, is one of the largest forest trees known.
1
Psidium guayava
and some species of Jambosa bear edible fruits; the former is especially valued.
Eugenia caryophyllata (Moluccas) is of economic importance, its unopened flower-
buds forming Cloves (Fig. 694). This tree is commonly cultivated in the tropics.
It has a beautiful appearance when the terminal groups of flowers (which in bud
634 BOTANY PART II

are of a bright red colour with four united, white petals ami when opened exhibit
4 j
numerous whitish stamens) are seen

through the shining foliage composed of

entire leaves. The corolla is thrown off
when the flower opens. In Fig. 694 the
inferiorovary, formed of two carpels, is also
seen in longitudinal section.
OFFICIAL. Eugenia caryophyllata
_ yields CAKYOPHYLLUM, Cloves. PlMEJfTA,
FIG. 693. Floral diagrams of Myrtaceae. A, Allspice, from Pimenla officinalis. OLEVM
.VvrfiM cammunis; li, Eugenia aromaiita. cAJAPUTl from Melaleuca leucadendron, a
(After EICHLER.)
tree Qf lesg height but resembling t h e

Eucalyptus trees ;
it is cultivated in the Moluccas (Burn) for the sake of the
oil it yields ;
its specific name refers to the white colour of the bark. OLEKM

Fio. 694. Eugenia faryofhyllnta (3 nat. size). Flowering -branch. A bud cut in half ami an opened
flower (aliout nat. size). OFFICIAL.

EUCALYPTI and EUCALYPTI ouMMi from Eucalyptus globuhis and other species.
Fi'.mily 8. Punicaceae. Single genus Punica. Punica granatum is a tree
SECT. TI PHANEROGAMIA 635

originally introduced from the East and now largely cultivated in the Mediterranean

Fio. 605. Punica granatum (J nat. size). 1, Branch bearing a flower and a bud. 2, Flower in
longitudinal section. 3, Fruit. (See text.) OFFICIAL.

region on account of acid refreshing fruits known as Pomegranates (Fig. 695).

its
Leaves small, entire. Flower with a stiff, red calyx, an indefinite number of
petals, and numerous stamens the 7-14 carpels
;

are arranged in two tiers, the upper of which

corresponds in number to the sepals, the lower
to the half of this (Fig. 696). Fruit enclosed
by a leathery pericarp with numerous seeds in
the loculi of both tiers. The external layers of
the seed-coat become succulent and form tin
edible portion of the fruit.
OFFICIAL. Punica granatum, from which c-
FIG. 096. Floral diagram of Punica,
GRANATI CORTEX IS obtained. granatum. (After EICHLER.)

Order 23. Umbelliflorae

Inflorescence as a rule an umbel. Flowers hermaphrodite,

actinomorphic, tetramerous or more usually pentamerous ; calyx
only slightly developed; a single whorl of stamens and an inferior
bilocular ovary, the upper surface of which forms the
nectary.
636 BOTANY PART II

Carpels two. A single pendulous ovule in each loculus. Seed with

abundant endosperm.

Family 1. Cornaceae. For the most part shrubs or small trees, rarely herbs.
Leaves simple, usually decussate. K4, C4, A4, G (2). Fruit, a berry or a drupe.
Cornus mas, the Cornelian Cherry (Fig. 697), expands its umbels of yello\\
(lowers before the leaves appear. Each umbel is subtended by four bracts. The
inflorescences for the succeeding year are already present in the axils of the leaves

by the time the fruit is ripe.

In Britain two species occur ;

C.samjiiiiK'n, the Dogwood, and

C. suecica, an arctic and alpine
plant which reaches its southern
limit in Germany.
Family 2. Araliaceae. Trees
or shrubs with alternate leaves,
which are often of considerable
sizeand simple, lobed, or com-
pound in shape. Floral formula,
Ky C5, A5,
; G(5). Flowers in
umbels or heads, often further
grouped in a panicle-like in-
florescence. The carpels vary
in number from 2-5 or more.
In Britain the only repre-
-> the family is the
'illative of

Ivy (Hcdera Helix}, a root-

elimber. The proper elliptical
leaf reappears on the
form
orthotropous shoots of older
plants, which in late summer
or autumn bear the flowers.
The leaves of the creeping or
FIG. 697. Cornus mas (J nat. size). 7, Flowering twig.
climbing plagiotropous shoots
.'.

Twig with fruits. Flower seen frcm above. 4, Flower

lobed and usually have
-.',
are
in longitudinal section. ('. !,. enlarged.)
shorter stalks (Fig. 698). Calyx
with five, pointed sepals corresponding to the five ribs on the inferior ovary. The
corolla is greenish in tint the large disc on the upper surface of the ovary attracts
;

the visits of bees. The fruits ripen during the winter and become blackish-blue
berries ;
these are eaten by birds and in this wav the seeds are distributed.

Family 3. Umbellifepae. Herbaceous plants sometimes of large

size. The stem, which has hollow internodes and enlarged
nodes,
bears alternate leaves these completely encircle the stem with their
;

sheathing base, which is often of large size. The leaves are only
rarely simple usually they;
are highly compound. Inflorescence
terminal, frequently overtopped by the next younger lateral shoot.
It is an umbel, or more frequently a compound umbel, the bracts

forming the involucre and partial involucres, or an involucre may be

wanting. Flowers white, greenish, or yellow other colours are ;
 Pro. Hull -r" lii'lix nat. size). 1, Erect flowering shoot. 2, Creeping shoot showing the
'
'''.is.
(i
distinct form of the leaves. 3, Fruits. A, Single fruit. 5, Flower from above. 6, Flower from
the side. 7, Flower cut through longitudinally. (4-7 enlarged.) PoiSOA'OUS.

Fie. 699. Umbelliferae, -Floral

diagram (N<7er).

!'!<;. 700. Fruits of l"nil>cllifrrni' in cross section. /. / <>

offlcii' A, < i.<i> I,,. .;, nium maculatum.

c,, /,.

Coriaii:- :>
(', nifxliticil nf'tcr a llgure by DRUDE).
637
 638 BOTANY

rare. K 5, C 5, A 5, G (2). The sepals are usually represented by

short teeth. The flowers at the circumference of the compound
umbel sometimes become zygomorphic by the enlargement of the
outwardly directed petals. Ovary always bi-carpellary and bilocular ;

KKI. 701.--Carum. carvi (J nat. size). In-

MuM-sci'iifc lM-aring fruits. Single flower,
and carpopliore bearing the mericarjts l-'in. 70-2. 1, Oenantltejutlulosa ( nat. size). 2. Group
(enlarged). OFFICIAL. of fruits. ,;,Single fruit (enlarged). Poisoxous.

in each loculus-a single ovule which hangs from the medium septum
with micropyle directed upwards and outwards. The upper surface
its
of the carpels is occupied by a swollen, nectar-secreting disc continu-

ing into the longer or shorter styles, which terminate in spherical

stigmas. Fruit a schizocarp, splitting in the plane of the septum into
two partial fruits or mericarps. In many cases the latter remain for a
time attached to the carpophore, which originates from the central
 PHANEROGAMIA 639

portion of the septum ; this separates from the rest of the septum and
bears the mericarps hanging from its upper forked end (Figs. 699-706).

For systematic purposes the fruits are of great importance. Each half of the
fruit has five ribs, beneath which the vascular bundles lie. The marginal ribs of
each partial fruit frequently lie close together at the septum or they may be
distinct they may resemble the three dorsal ribs or differ more or less from them.
;

Between the five primary ribs four secondary ribs are sometimes present. Usually
furrows (valleculse) occur between the ridges, and beneath each furrow a large oil
duct (vitta) is found, extending the whole length of the fruit. On either side of

Flo. 703. Slum latifolium (J nat. size). Poisonous.

the carpophore a similar oil duct is present in the septum, so that each mericarn
has six of these vittse (Fig. 700). In some species additional small ducts are
present. The form of the fruit as seen in a cross section differs according to
whether the diameter is greater in the plane of the septum or at right angles to
this. The character of the marginal and dorsal ridges and the presence or
absence of secondary ridges or vittae serve to distinguish the fruits, and are
indispensable aids in determining the species. Since many of the fruits are
employed in medicine or as spices, while others are poisonous, their distinction
becomes a matter of importance. The endosperm of the seeds contains a oil fatty
as reserve material.
The most recent revision of the family is by Drude in
En<*ler-Prantl Nat
/;//"// :cnfam. ; an older useful division is founded on the
shape of the endosperm.
1. Orthospermae.The endosperm flat or; slightly convex on the^ventral side
640 BOTANY PART II

i.e. on the side towards the plane of junction of the two mericarps (Fig. 700, 1, 2).
The majority of the species belong to this group.
///</;(//// is a civcpin;;
plant growing in marshy places with entire, peltate leaves and simple umbel*.
Sanicula and Astrantia have simple umbels, which in the latter are surrounded
by a petaloid involucre. In Eryngium the plant is
usually spiny and bears

Fie. 704. Cicuta rirosa. Rhizome cut through longitudinal!} (A nat. si/r). Fruit
(enlarged). Poisosocs. 1

simple umbels, which approach closely to being capitula the involucral leaves an-
;

often coloured. Bupleurum has entire leaves. PimpincUa, Burnet-Saxifr;ig .

/'. Anisum, Anise, an annual plant, the seedlings of which exhibit increasing
is

subdivision of the lamina in successive leaves. Carum C'crri, Carroway. lias

long been cultivated (Fig. 701) leaves bipinnate, the lowest pinnae membling
;

stipules. The large lower pinnules are usually placed horizontally on tin- vntii-al
rhachis of the leaf; the terminal pinnules are simple and linear. The termina-
 PHANEROGAMIA 611

umbel, the flowers of which open first, is overtopped by the lateral umbels arising
from the leaf-axils. Biennial. Carum Bulbocastmmm has a perennial, tuberous
rhizome its seedlings have only one cotyledon.
;
Foeniculum (Fennel) and
Levisticum (Lovage) have yellow flowers. Petroselinum (Parsley), Pastinaca
(Parsnip), Daucus (Carrot), Aplum (Celery), and Anethum (Dill), are used as

vegetables. Cicuta (Water-Hemlock, Fig. 704), Sium (Water-Parsnip, Fig. 703),

Oenanthe (Fig. 702), and Berula, are marsh- or water-plants. Aethusa cynapium
(Fool's Parsley, Fig. 705) has the ribs of the fruit keeled umbels with three ;

Kii;. 705. Aethusa cynapium (5 nat. size). B, Single umbeL '', Fruit (enlarged). Poisoxocx.

elongated, linear, involucral leaves directed outwards. All the last-named plants
are poisonous. Archangelica ojficinalis is a conspicuous plant reaching a height of
2 metres, with large, bipinnate leaves provided with saccate, sheathing bases the ;

greenish flowers are markedly protandrous, the styles and stigmas only developing
after the stamens have fallen. In Carum, Oenanthe, Cicuta, and Heracleum, and
notably in the large species of Ferula, the flowers become polygamous or of
separate sexes by abortion of stamens or carpels ; sometimes the plants become
dioecious. The ovary i.s completely wauling in the male flowers.
2. Campylospermac. The ventral side of the endosperm is traversed by a

longitudinal groove (Fig. 700, 3). Scandix, Anthriscus (Beaked Parsley), Ohaero-
2 T
642 BOTANY

I (Chervil). GW
//'/;/ uxx-i/liit u,,i. the Hemlock, is a biennial
plant often of
considerable height ; it is completely glabrous, the stem and leaf-stalks often with
purple spots ;
leaves dull green, bi- to tri-pinnate. The ultimate segments end in
a small, colourless, bristle-like tip. Fruit with wavy, crenate ridges and without oil-
ducts in the valleculte. The whole plant has a peculiar, unpleasant odour (Fig. 706).

FIG. TOti. Conium maau.la.ttuH. (J nat. size). UrnciM. ami I'uisuxucn.

3. Coclospcrmac. The ventral side of the endosperm is concave (Fig. 700, 4).
Coriandrum sativum is an annual plant flowers zygomorphic owing to the enlarge-
;

ment of the sepals and petals at the periphery of the umbel. Fruit spherical ;

mericarps closely united, with ill-marked primary ridges and somewhat more
distinct secondary ridges.
OFFICIAL. Conium maculatum yields coxn FOLIA and CONII FRUOTUS. Ferula
foelida (Persia), ASAFETIDA. F. yalbaniflua and other species (Persia), GALBANUM.
 PHANEROGAMIA 643

Dorciiin Aiiiiiwniaeuiii (Persia), AMMONIACUM. Pimpinella anisum, FRUCTUS ANISI.

CoriandriiTH sativum, CORIANDRI FRUCTUS. Foeniculum capillaceum, FOENICULI
FRUCTUS. Carum carvi, CARUI FKUCTUS. Anethum (Peucedanu'in) yraveolens,
ANETHI FRUCTUS. Fcritla SUIflbul, SUJIBUL RADIX.

Order 24. Hysterophyta

This order provisionally groups together a number of plants which are for the

Flu. ~V~.Ai-ixt<il<-n-lii<t <:leiiMtiii* (\ n;it. si/.c). To the right a tlovvcr cut through longitudinally.

most part parasitic. The several families show no close relationship to one another
or to other families.

Family 1. Aristolochiaceae. Non-parasitic herbs and climbing shrubs with

644 BOTANY

simple, cordate leaves. Flowers with simple, gamophyllous perianth. Androeciurn

and gynaeceum united to form a gynostemiuin. P3, A64-6, (Mi Interior. .

septate ovary. Fruit a capsule.

Asarum eurojmeum occurs in Britain ;
flowers actinomorphic. Aristufucliin
clematitis (Fig. 707) has sygomorphic, protogynous flowers whieli are pollinated

by small insects (cf. Fig. 243, p. 312). An erect herb with large, alternate' leaves,
bearing the numerous flowers in their axils.
Family '1. Santalaceae C
:(7
). Green plants growing in the soil and parasitic

Fin. 70s. !"/>< i 'ill, inn. With ttowris and fruits, (i nut. si/r.)

on the roots of other plants from which their haustoria obtain nutrient materials.
In Britain, Thesium.
OFFICIAL. Santalum album, the wood of which when distilled yields OLEUM
SA.NTALI. The wood is also of economic value.
Leafy, parasitic shrubs, living on the branches of
:i8
Family 3. Loranthaceae ). (

trees. They are most abundant in the tropics, and, for instance in South America,
add to the beauty of the forest by their brightly coloured flowers.
r
Loranthus europaeus, on Oaks in Europe. In Britain l iscum album(ig. 708),
the Mistletoe, occurs as an evergreen parasite on a number of trees. It has opposite,
obovate leaves. Stem swollen at the nodes. The inconspicuous flowers open in
early spring they are diclinous and dioecious, usually bimerous, and are borne in
;

five-flowered dichasia. The white berries ripen during the winter (December to
March) and are eaten by birds. By means of a viscid layer of the periearp they
 PHANEROGAMIA 645

adhere, together with the seeds, to branches against which the birds rub
their beaks. The root on germination forms an expanded, attaching disc. From
the centre of this a sucker, which has no root-cap, penetrates the rind of the
branch as wood. From the base of this lateral roots arise, which grow
far as the
in the rind giving off secondary suckers. As the wood of the host grows in
thickness year by year, the tips of the suckers become embedded in the new
wood. Their tips become converted into permanent tissue, while further growth
in length is effected by a zone corresponding in position to the cambium of the
host plant.
4. Rafflesiaceae M 5. Balanophoraceae
4U
The plants be-
Family ( ). Family ( ).

longing to these families are almost all tropical. They are parasitic, without chloro-
phyll, and with their vegetative organs 1'educed to haustoria. They have sometimes
very large flowers (Rafflesia] or large inflorescences bearing numerous small flowers.
Rajflesia Arnoldi (Sumatra) has the largest of all known flowers, which attain a
diameter of 1metre. Many species of Balanophora are apogamous.

15
Series II. Sympetalae ( )

Perianth differentiated into calyx and corolla. Corolla gamo-

petalous. Flowers usually pentamerous, always cyclic. Two groups
of Sympetalae can be distinguished. A. Pentacydicae. Flowers
with five regularly alternating whorls. K5, C(5), 5 + 5, G(5). A
B. Tetracydicae. Flowers, by omission of the inner whorl of
stamens, no trace of which persists, composed of four regularly alter-
nating whorls. Ko, C(5), A 5, G(5) or more commonly G(2) by
reduction in the number of carpels.

A. PENTACYCLICAE
41
Order \.
Erieinae( )

Plants with actinomorphic flowers; androecium obdiplostemonous ;

stamens not adherent to the corolla. "

Pollen-sacs with an exothe-
"
cium pollen-grains frequently cohering
;
in tetrads. Ovary, as a
rule superior, multilocular.
Family 1. Ericaceae. Evergreen, shrubby
plants with small, often needle-shaped leaves.
Anthers opening by pores or splits, frequently
provided with horn-like appendages, on which
account the termed Bicornes.
group is also
Fruit, a capsule, berry, or drupe.
Seeds with
a well-developed embryo and abundant endo-
sperm (Figs. 709, 710).
FIG. "Oi>. Kloral (liairiani of
IMPORTANT SUB-FAMILIES AND GBNKBA. A. With j /;, (Ei-irac-a.-).
,i
Miperior ovary 1. Ericeue.
:
K4, 0(4), A4 + 4, G(4).
Corolla persistent. Erica, calyx shorter than the corolla, includes some British
species (E. Tetrulix) and many native to the Mediterranean region and the
Cape. Calluna, calyx longer than the corolla. C. vulgaris, Heather or Ling,
 646 BOTANY PART II

occurs on both dry ami wet soils in western Europe. It ascends into
alpine regions, and by itself or in association with a tew other plants rover.-
extensive tracts of country known as heaths or moors. '2. Ar)>nti<\ Flowers
pentamerous. Corolla deciduous. Aiiilriniil<i occurs on moors in the northern

hemisphere fruit, a capsule.

; Afi-tnn/c/i/iytos Uva m-si. ciiviini]>olar. in similar
localities, has small, entire, leathery leaves. Fruit, a live-seeded drupe i_ 1

710). 3. Rhodorcac. Corolla deciduous. Capsule septioidaL ]!1<<xlodendron,

Alpine Rose. There are a few European species many t'orins. especially Asiatic ;

ornamental sliruhs.
species, are cultivated as Lalum //////..// is a poisonous, evea
green shrub, occurring on moors in Europe leaves rolled round at the DUrgini : ;

beautiful white umbels of flowers. Corolla polypetalous, star-shaped.

B. Ovary inferior: Voceincae. Vaccinium myrtillus, Blaeberry. i> a 1\\ .

r
deciduous, shrubby plant. l ritis idaea, Cowberry, is evergreen, and together
.

with the preceding species is found in hilly regions.

FIG. 710. Arctostiijihiilos I'm tirsi. 1, Flowering branch ; .'. Mowers in longitudinal M-ction :
9,
r fruit in transverse section. BERC ;m<l
ixillen-j^-ains ; I,, fruit ; .
>, (After SCHMIHT.)

OFFICIAL. Arctostajihylos UTU in-*i yields VAK n:.si KUI.IA. i

Family 2. Perennial, sometimes evergreen, herbsoaoUB plants.

Pyrolaceae.
Corolla often polypetalous. Seeds very small embryo composed of only a few ;

cells. Pyrohi. several British species, with the flowers solitary or in racemes.
Monotropaw a saprophyte devoid of chlorophyll. M. hyjwpitys, Bird's Nest, occurs
rarely in woods, has a yellowish, succulent stem, bearing scale leaves below and
ending in a racemose inflorescence with a few flowers.

Order 2. Diospyrinae
Actinomorphic, usually pentamerous flowers with a diplostenionous androecium.
Antisepalous stamens frequently reduced. The Sapotaceae is a tropical family ;

the plants contain latex. Species of Palaquium and Payena from the Malayan
Archipelago are the trees from which gutta-percha i.> obtained. Flowers hexa-
merous (Fig. 711). Balata is obtained from Mini usu/i* trees found throughout ;

the tropics.
Ebenaceae. Diimpyros Kuki is a Japanese fruit tree ; ]>. W>< HH,,I. ebony.
Styracaceae. The origin of Benzoin
(BENZOIXUM), an official resin, from
Styrax Benzoin, though generally assumed, is open to doubt.
 PHANEROGAMIA 647

FKI. 711. Palaqnimn (lutta. (i nut. sixc. At'tt-i- S< in MANN and A. MEYER.)

Order 3. Primulinae

Flowers sympetalous, actinomorphic with only one whorl of

stamens the latter are antipetalous, and are adherent to the corolla.
;

Ovary unilocular, superior, with a free-central placenta. Ko, C(5),

A5, G(5).

Family 1. Primulaceae. Jin I

is, often with Conspicuous flowers. Calyx green.
Style, unbranched. Numerous ovules. Fruit, a capsule (Figs. 712-7H).
648 BOTANY PART II

IMPORTANT GENEKA. /V/< "/, with several British species. Herbs with rosettes
of leaves. Flowers solitary or in umbellate inflorescence* The capsules open by
five teeth at the summit. Commonly grown as ornamental plants. Anagallis,
capsule opens by a lid (pyxidium). Stem prostrate with leaves in decussate pairs
f
Fig. 714). Lysimachia, large herbaceous plants, with erect or creeping stems and

Fio. 712. PrflnuUceae. Floral

diagram (I'nmulc.).

Fit;. 714. AwmaUitarveitthQ nat. si/i-).

FK;. 713.- I'li'-hiim a .

in-iijHifiiiii. A. entire plant : />'. fruit. Longitudinal section of (lower ami
capsule at ilehiscence (cular^fd).

4-
decussate leaves. Cyclnmi'ii flowers bent back on long stalks
i
, tips of the ;

petals bent back. .Many species and hybrids are in cultivation. C. europaeum, the
Alpine Violet (Fig. 713), is a herbaceous plant found in the Alps the somewhat ;

fleshy leaves have long stalks, and are borne on a large perennial tuber, which
originates from the hypocotyl of the seedling. The stalks of the fruits become
spirally rolled.
Poisoxors. The tubers of Ci/cfa num 1'urojnn ///.
though harmless when cooked,
are poisonous in the fresh state. Anagallis is slightly poisonous. Primula obconica
 PHAXEROGAMIA 649

(and P. si'iie'iisis) secrete in their glandular hairs certain substances which fre-

quently give rise to rashes and to inflammation of the eyes of those handling the
plants.
Family 2. Plumbaginaceae. Herbs with a dry, membranous, but often brightly
coloured calyx. Style branched. The ovary contains a single ovule. Species of
Armeria and Statice are common plants of salt marshes.

B. TETRACYCLICAE
Order 4. Contortae

Plants with decussate, usually simple leaves and actinomorphic

flowers, the corolla of which is often contorted in the bud. Stamens
epipetalous. Ovary of two carpels, superior.
Family 1. Oleaeeae. Shrubs or trees with-
out stipules. Flowers tetramerous, but with
only two stamens. K4, C(4), A2, G(2).
Corolla with valvular or imbricate .-estivation.
Ovary bilocular; two ovules in each loculus

(Fig. 715).

IMPORTANT GENERA. Oh i-m-ninn the Olive f

43
), a
FIG. 715. Oleaceac. Floral
native or the Mediterranean region and ot western Asia,
dia"ram(Sj/-/''/").
where also it is cultivated. It is a tree with a poorly
developed crown of slender pendulous branches bearing lanceolate, simple, entire
leaves, which are dark green above, greyish green beneath. The (lowers are borne
in axillary racemes or panicles. The fruit is a drupe, both the succulent exocarp
and the endosperm of which contain a fatty oil (Figs. 717-719). Ligustrum vulgare,
the Privet. Jasurinnm and Kiirin<j<i (Fig. 715) are grown as flowering shrubs.
/'i-n.i'i'iiiix. the Ash, has
inipari|iinnate leaves: F. excelsior has apetalous, anemo-
philous flowers, \\hieh appear before the leaves. /'. ariius, the
Flowering Ash. has
a double perianth and is entomophilous it is polygamous, having hermaphrodite
;

flowers as well as female flowers with black, sterile anthers the corolla is divided ;

to the base (Fig. 716).

OFFICIAL. Olca <:urojn'n yields OLKI'M IH.IVAK.
Family 2. Loganiaceae. Herbs, shrubs, or trees with stipulate leaves. K f>,
0(5), A5, G(2). Tropical. Species of X/,-,/i;/inos, which are trees or lianes climbing
liy means of hook-tendrils, yield the arrow poison
as well as the well-known Curare
of South America, and that used by the Malays.
OFFICIAL. Stri/r/nmx //./ /(/////><> is a small tree or shrub of southern Asia, the
fruits of which are berries with a firm rind in the succulent pulp a small number
;

of erect, circular, disc-shaped seeds are embedded (Fig. 720). It yields xrx
VOMICA and STKYCIIXIXA. Cl'.I.M-'.Mll RADIX is obtained from <!<h<'ni inm iliil nl,. ii ii

which is a native of the south of North America.

Family 3. Gentianaeeae. Herbs, with exstipulate leaves.

Corolla contorted in the bud (Fig. 721). The two carpels are united
to form a unilocular ovary. Ovules numerous.
41
i,
i> a genus with numerous species. Plants of
larger or smaller si/e. especially abundant in the Alps.. Flowers brightly coloured.
 .-

Fio. 710. Frojfinvx amim (A nat. si/e). Flower and fruit.

Kic. 717. Olec eiirojmit. A, Corolla spread out B,

Calyx and ovary in longitudinal section. (Fn-
larged. Aft>r ESCI.KR-PRAXTL.) l-'i... :is. Q Drape. A.

650
SECT. II PHANEROOAMIA 651

This genus affords one of the best examples <>! si-a>on;il dimorphism, i.e. the split-
ting of a species into two closely related forms which develop at different season.-.
Since the height of the vegetative period of the Alpine meadows coincides with
their annual this expresses itself in the distinction of an early form, fruiting
mowing,
before the meadows
are cut, and a late form developing after this has taken place.
Eri/thraea, Centaury (Fig. 722), anthers spirally twisted. Menyanthes, Bog-Bean,
with alternate, trifoliolate leavt->
and long-stalked racemes of white
flowers with hairy petals. Lim-
nanthcmum, aquatic plants with
floating leaves.
OFFICIAL. Gentiana lutca arid
other species yield GENTIANAE
KADIX. CHIKATA is obtained
from Su'ertia chirata (N. India).

Family 4. Apocynaeeae.
Evergreen herbs, shrubs
or trees, without stipules.
Especially numerous in the
tropics. Laticiferous cells
in all parts of the plant. K5,
0(5), A5, G(2). Petals con-
torted in the bud. Stigma ring-
shaped. Carpels only united
in the region of the style,
free below (Fig. 723) and
separating after fertilisation.
Usually two follicles. Seeds
provided with a tuft of hairs.
IMPORTANT GENERA. The only
British species is Vinca minor,
the evergreen Periwinkle, occur-
ring in woods (Fig. 726). Nerium
oleander (Fig. 725), a native of S.
Europe, is commonly cultivated.
It has greyish -green, lanceolate
leaves which are decussate or in Km. 71!'. Olea evropea in fruit. (J nat. size.)

whorls of three ;
flowers rose-

coloured, sweet-scented and conspicuous. The whole plant is extremely poisonous.

Wa
OFFICIAL. Strop/tanthus /combe and S'. hispidus ( ) (Fig. 724), lianes of tropical
Africa, yield STKOPHAMHI SEMINA.
43
Caoutchouc ( ) is obtained from Kickxin ehisticn and other species, trees of

tropical "W. Africa. It is numerous species of Landolphia

also obtained from
\L. Kirkii, Heudelotii, comvrensis, etc.), lianes which are, or formerly were, common

throughout tropical Africa. Together with species of Carpodimtu (Angola) which

yield the root-rubber, these plants supply the main part of the rubber obtained
from Africa. Hancornia speciosa, a tree of the dry Brazilian Campos to the south
of the Amazon " "
region, yields the mangabeira rubber. Jl'-illomjlibfio firma,
652 1JOTANY PART II

W. flavescens, and other species of this Malayan genus of liaues are also rubber-

Kic. 7.0. Strychnos nux voniirn (\ nat. size). Fruit and seed whole
and in cross section. OFFICIAL and Poisoxovs.

FIG. 721. Gentiaita lutca. and h, Kl(i\vci-linds(n:it. sixr). sliowinj;

calyx () and twisted corolla (/<) c, transverse section of ovary.
; KII^. T'J-J. F.rythraea cen-
OFFICIAL. (After BERI^ and SI-HMIDT.) tin: ,-iii in. (i nat. size.)

yielding plants. Gutta-percha is present in the latex of Tnh, -mx iitmttuna Donnell
,

Smithii, which is used as a shade-tree on coffee plantations in Central America.

 PHANEROGAMIA 653

Since, hitherto, only some Sapotaceous plants,

which are difficult to cultivate, have yielded
gutta, this plant will soon take an important
place in the supply.

Family 5. Aselepiadaeeae. Similar

and closely related to the Apocynaceae.
Corolla contorted in the bud carpels ;

free,only united by the prismatic stigma ;

latex in all the organs. Stamens united
at the base with dorsal, FIG. 723. strophanthw hispid, ovary
nectar-secreting
in longitudinal section. after
appendages forming a corona. The pollen
ENGLER-PRAXTL.)

Fio. ~24.Str<>p1iitiitliii.-< liisjiiilii* (i nat. size. After MEYER and S< HI-MANN). Fruit (I nat. si/..).

Seed(Jnat. si/c). (After SCHUMANN in EXGLEB-PBAXTI..)

 54 BOTANY i'AKT II

of each pollen-sac is united into a pollinium, the stalk of which is

attached to a glandular swelling (adhesive disc) of the angular
stigma.

These adhesivr discs alternate with the stamen- so that the two jiollinia attached
to each disc belong to the halves of two adjoining stamens. Visiting insects
iviiiove, as in the Orehidaceae, the jiollinia and rarry them to another flower

(Fig. 727).
IMI-OKTAM (IKXKKA. Vincetoxicum ojficiiuile (Fig. 7->) is a European herb
 PHANEROGAMIA 655

Fi<;. 728. Vincetoxicum officinale

(i nat. size). Poisoxocs.

7 lit I. Vinca minor ($ nat. si/.p).

c. T'JT. Asdepias cuntssavica. A, Flower; an, ;ui<h<vimii (x 4); II, calyx and ^yimt'cr
/, ovary fr, oorpuscula ( x
; (5) C, pollinia (more highly iiiii-nified).
; (After BAII.UIS.)
 65 BOTANY

with inconspicuous, white (lowers ami hairy seeds which are borue in follicles ;

poisonous. Other forms are mostly tropical or suh tropical. The succulent -

species of Stapelia, Hoodia, Tnchocuiilmi. etc.. which resemble Cactaccac in

habit, and inhabit S. African deserts, and IHsi-linlin r<irili-xi<ni c"
;

). the peculiar

pitcher plant of the Malayan region, deserve special mention. IIim <-nriLosa is

frequently cultivated.
OFFICIAL. Hemidesmus iiidictts yields HKMIDF.SMI IIAIUX.

Order 5. Tubiflorae

Flowers pentamerous, actinoraorphic or zygornorphic. Carpels L'.

Ovary superior, bilocular, with two ovules, which are frequently

separated by a false septum, in each loculus.

KK.. 7_".'. i'<irnli-nhix //Hsix (i nat. Pro. 780. JBwjoniitiH jmtfo (J nat size. After.

Longitudinal section of the BKR<; and SCHMIDT). OFFICIAL.

size).
ttoxxer and a capsule (enlarged).

Family1. Convolvulaeeae. Herbs or shrubs, less commonly

trees, with alternate, sagittate or cordate leaves. The majority are
twining plants. Many contain latex. Corolla widely funnel-shaped,
folded and twisted
only slightly lobed ; in the bud it is longitudinally
to the right. Stamens five. Ovules basal in the two or four cavities
of the ovary. Fruit, a capsule.
IMPOUTAXT GKXEKA. Convolvulus arvennt (Fig. 729), a perennial, twining,
herbaceous plant occurring everywhere by waysides, and as a weed in corn-fields.
 PHANEROGAMIA 657

Flowers solitary, long-stalked, situated in the axils of the leaves and sometimes in
the axils of the bracteoles of another flower. Calystegia has two large bracteoles
placed immediately beneath the calyx. C. sepium. The Dodder (Cuscuta) ( 47 ) is a
slender parasitic plant containing very little chlorophyll, which attaches itself by
means of haustoria to a number of different host plants. The small flowers are
borne in dense clusters (cf. p. 229, Fig. 202). Ipomoea is for the most part a
tropical genus ; several species are cultivated as ornamental plants. /. jjescaprae
is one of the strand plants of tropical countries.
OFFICIAL. JALAPA is obtained from Exogonium purga (Fig. 730), a twining

FKI. 732. Kchluin vulgare. Inflorescence (J nat. size),

flower and fruit, composed of four nutlets (enlarged).

/ \
Kid. 731. Borago officinalis. a, FIG. 733. Floral diagrams of Verbena officinalis (A), (after
Flower ;
b and c, fruit (nat. size). BICHLER), and of Lamiiim (Labiatae) B.

plant, with tuberous lateral roots, occurring on the wooded, eastern slopes of the
Mexican table-land. SCAMMONIAE RADIX is the dried root of Convolvulus Scumiinmin
(Asia Minor).
Family 2. Polemoniaceae. Carpels usually three. No milky juice. Polemonium
coeruleum occurs in Britain. Cobaea seandens and Phlox are cultivated.
Family 3. Hydrophyllaceae. K 5, C (5), A 5, G (2). Inflorescence a
cincinnus.
Family 4. Boraginaceae. Herbs with alternate leaves. These and the stems
are covered with coarse hairs (Asperifoliaceae). Inflorescence always cincinnal ( 48 ).
Flowers actiuomorphic or zygomorphic. Petals frequently provided with scales
2 IT
658 BOTANY

.standing in the throat of the corolla.

Ovary always bilocular but divided by
false septa into four one-seeded nutlets.
The style springs from the midst of the
four-lobed ovary. Ovules pendulous.
Fruit, a schizocarp, separating into four
portions (Figs. 731, 732).
GENERA. Symphytum (Comfrey),
Borcujo (Borage), Anchusa (Alkanet),
Echium (Bugloss), Myosotis (Forget-me-
not), are among the commonest and most
conspicuous herbaceous plants of our flora ;

allhave entire, alternate leaves, covered

with harsh hairs and relatively large
Mowers ofa lighter or darker blue, grouped
in complicated inflorescences. InPulmon-
aria officinalis, which flowers in spring,
the colour of the corolla changes from red
to a bluish violet. Cerinthe is completely
glabrous, and has a waxy covering to the

epidermis. Heliotropium, a favourite

ornamental plant, has an undivided
ovary.
Family 5. Verbenaceae. Shrubs or
trees, rarely herbs, with zygomorphic
flowers. K f, C (5), A 4, G (2). Distin-

Fl<;. 734. '.'"/"/''-' <-hi-iJi-ii'-ii. O, HUN-IT; li,

tin- same with calx x remove,) ; <. cnri.lla cut

(P[M-H. slmwini: stamens ami style; d, calyx
ami -\nai-i-i-iuii: r. fruit. fflciM.. (a, 6,
init. sixc : i; <>. < x 2.)
Flo. 785. Lanupfula vern (A nut.
OFFICIAL.
 PHANEROGAMIA 659

guished from the following family by the terminal position of the style (Fig.
733 A). Only Verbena officinalis occurs in Britain.
Lantana is a widely spread weed in the tropics.
Tectona gratidis ( 49 ), a deciduous tree of the dry
regions of the East Indies, yields the important
timber TEAK. Avicennia ( 80), trees of the mangrove
vegetation, with viviparous fruits and pneumato-
phores (Fig. 213). Clerodendron ( 51 ) includes some
myrmecophilous species.

Family Labiatae.
6. Herbs or shrubs
with quadrangular stems and decussate
leaves without stipules. Leaves simple ;
plants often aromatic owing to the presence
of glandular hairs. Flowers solitary in the
axils of the leaves, or forming apparent
whorls. The small inflorescences are di-
chasia or double cincinni, and are often
united in larger, spike- or capitulum-like
inflorescences. Flower zygomorphic (Fig.
734). Calyx, gamosepalous, with five teeth;
corolla two-lipped, the upper lip consisting
of two, the lower of three petals ; stamens
in two pairs, two long and two short, rarely
only two. Ovary of two carpels, each of
which deeply indented in the plane of
is

the false septum. Style springing from

between the lobes ; stigma bifid. A ring-
shaped nectary situated at the base of the
ovary. Fruit, a schizocarp, consisting of
four nutlets.

IMPORTANT GENERA.
The Labiatae include a
considerable proportion
of our commonest native
spring and summer
flowers ; Lamium, Gale-
opsis, and Stachys have
the upper lip helmet-
shaped, Ajuga has it very
short, while in Teucrium
the upper lip is deeply
divided. Nepeta and
Glechoma differ from the *f
majority the order,
of
in having the posterior
"
*'"' 3li - s "'
stamens longer than officinalis. Flowering shoot (J nat. size). Tubular
" is " lay
l "" 8tameBa (eillar ed
those of the anterior pair. >-

Salvia, Sage (), is a genus containing numerous species ;

the flowers are protaudrous
660 HOTANY

and are characterised by the suppression of two of the stamens usually present.
(Fig. 736). The two fertile stamens have a very long connective, with the anterior
theca, in which pollen is formed, protected beneath the upper lip. The posterior
theca, the development of which differs in tin- species, projects into the entrance of
the corolla tube leading to the nectary (cf. p. 312, Fig. 244). A
bee visiting the
flower, presses this theca up-
wiinlsand thus bringsdown the
other arm of the lever, formed
by the elongated connective,
upon its back. In older flowers
the bifid stigma occupies such a
position that it will rub olf pol-
len brought by an insect from
another flower. Many species
of Salvia are cultivated as or-
namental plants, and are often
conspicuous, rather by reason
of their brightly coloured
bracts than by their flowers.
Labiatae are especially abund-
ant in the xerophytic forma-
tion of shrubby plants in tin-

Mediterranean region to which

the name Maquis is given :

Salvia officinal IK, M<irrnliinni,

Phlomis, and species of Li
dula. Rosmarinus
is a showy shrub with small
narrow leaves revolute at the
margin, and bright blue flowers
with only two fertile stamens.
Many species are grown as
kitchen herbs, e.g. Origanum
majorana, Satitn'jn hurti-nsis.
Ocimum basilicum, Salvia and
Thymus. Thymus vulgaris
from the Mediterranean, like
our native T. serpyllwm, is a
low, shrubby plant with poly-
gamous flowers ;
there are

larger.hermaphrodite and small,

purely female flowers. Some
FIG. 737. Melissa officinal ix (i nat. size.
species ofMentha are similar.
After liERi; ;in<l SCHMIDT).
The widely cultivated .'/.
/"/
-

hybrid between M. ririttis and M. aquatica it is completely sterile, and has

rita is a ;

been propagated by runners from ancient times. 31. crispa, another commonly culti-

vated form, is a crispate variety of M. viridis M. longifoUn, var. imdulntn. M.

;

sylvcsfris, etc.
OFFICIAL. officinalis yields OLEUM ROSMARIXI.
Rosmarinus Lin-amlnln ttn
(Mediterranean region), OLEUM LAVAXDM.AE. .!/////<" />i/im'i'i. 01 .KI M MKXTHAE
1'ii'EitiTAE. M. viridix, oi.KfM MEXTHAE viuiins. M. Ofventit and .17.
yield MENTHOL. Thymus vulgaris and Monarda punclata yield THYMOL.
 PHANEROGAMIA 661

Order 6. Personatae
The Personatae are connected by the small family Nolanaceae to
the Convolvulaceae, from which the Tubiflorae are also derived. The
flowers are actinomorphic or zygomorphic. Their typical form has
also the floral formula K5, C(5), A 5, G(2). There are, however,
no false septa in the ovary and the number of ovules is usually a
larger one.
Family 1. Solanaceae. Herbs or small woody plants, with alter-

nate, exstipulate leaves, and nearly always actinomorphic flowers,

K5, C(5), A5, G(2). Corolla expanded or tubular; petals plaited
in the bud. Ovary bilocular, septum inclined obliquely to the
median plane. Ovules numerous, on a thick placenta. Fruit, a
capsule or a berry. Seeds with endosperm ; embryo usually curved
(Fig. 738). In many Solanaceae, as will be
further referred to in the special descriptions
below, the inflorescences exhibit apparent, extra-
axillary branches and paired leaves. Anatomi-
cally the order is characterised by possessing
bicollateral vascular bundles.
IMPORTANT GENEKA AND SPECIES. () Fruit, a
berry The Deadly Nightshade (Atropa Belladonna, Fig.
:

739) is a perennial herb of shrubby habit, springing

from an underground rhizome. It is a native of Europe FIG. 738. Solanaceae. Floral
and western Asia, occurring in less dense woods. The diagram (Petunia).
shoots are, to begin with, orthotropous and radial, and
bear alternate leaves and a terminal flower, which only rarely produces fruit.
Below this terminal flower branching and the development of leaves commences
in. as a rule, three,
equally vigorous, lateral shoots. Each lateral branch with its
further cincinnal branching forks and assumes a dorsiventral habit. By the pre-
dominance of one of the two axillary shoots and the carrying up of the subtending
bract upon it, the terminal flower at each grade of branching becomes apparently
axillary. The large, subtending bract is borne up beside the smaller one belonging
to the next higher axillary bud, which is usually undeveloped, so that the leaves

appear to be borne in pairs. Flower with a short, wide, tubular corolla of a dirty
purple colour. Calyx enlarging after fertilisation beneath the bluish-black fruit ;

the position of the obliquely placed septum of the latter is recognisable externally
by the presence of a shallow groove.
Many species of Solanum occur as weeds. Flowers actinomorphic. S. niyrum,
Nightshade S. dulcamara, Bitter-sweet (Fig. 740), is a shrubby plant, climbing by
;

HUM us of its stems and petioles, and especially common in thickets by the banks
of streams and similar situations. S. tuberosum, the Potato. Lycopcrsicum, the
Tomato. Capsicum annuum, Spanish Pepper, has a dry, berry-like fruit. These
plants resemble Atropa in their branching and the position of their leaves.
/ Fruit a capsule Datura Stramonium, Thorn-apple (Fig. -741), is an annual
:

plant, widely spread in Europe, Asia, and N. America. It has incised, palmately-
veined leaves and large, white, terminal flowers. The spiny fruits split at the
summit into four valves. The first terminal flower is developed early, and the
plant then exhibits profuse dichasial branching. Since the subtending leaves are
662 BOTANY PART II

adherent to the axillary shoots, which as a rule develop equally, tin- branching
appears to be extra-axillary, a terminal flower or fruit being situated in the t'nik.

FIG. 739. Atropa Belladonna (J nat. size). Orrn IM. ami /'o/.v/.voj .-.

Nicotiana tabacum (Figs. 742, 743) is a South American plant with numerous
cultivated varieties, which are grown both in Kuropc and the tropics. Its large
alternate leaves, which bear numerous glandular hairs, form, after being dried and
SECT. II PHANEROGAMIA 663

prepared, TOBACCO. Flowers elongated and tubular, borne in terminal panicles.

Hyoscyamus niger, the Henbane (Fig. 744), is an annual plant occurring in Central
Europe, North Africa, and Western Asia. The radical leaves have long stalks,
those of the flowering stem are sessile they are all clothed with glandular hairs.
;

Flowers slightly zygomorphic, of a dull yellowish - violet colour with darker

markings ; inflorescence, a cinciimus. Fruit, a pyxidium. Petunia and Salpiglossis
are favouritegarden plants with obliquely zygomorphic flowers.
All Solanaceae are more or less poisonous partly on account of the presence of

Fio. 740. Snlunitm dulcamara (k nat. size). Poisonous.

considerable amounts of alkaloids or poisonous glucosides. Species of Solanum,

Atropa, Datura, Hyoscyamus, and Nicotiana are among the most poisonous plants
met with in this country.
OFFICIAL. Capsicum minimum yields CAPSICI FRUCTUS, Atropa belladonna
yields BELLADONNAE FOLIA, BELLADONNAS RADIX and ATROPINA. Datura
Stramonium, STRAMONII SEMINA and STRAMONII FOLIA. Hyoscyamus niger,
HYOSCYAMI FOLIA.

Family 2. Serophulariaeeae. Herbs, with opposite or alternate,

exstipulate leaves. Flowers zygomorphic. Corolla not plaited in
the bud. Number of stamens often incomplete. Carpels median
(Fig. 745). Fruit, a bilocular capsule.
 664 BOTANY

IMPORTANT SUB-FAMIUES AND OKSEUA. (a) Antirrhincue, not parasitic.

FIG. "41. Datura Stramonium (J uat. sixc).Mature fruit after doliiscf IIQT. OFFICIAL
and VOISOKOVS.

Vcrbascum (Fig. 746), the Mullein biennial herbs, which in the first season form
;

a large rosette of leaves from which the erect inflorescence arises in tiio second
 PHANEROGAMIA 665

year. The inflorescence sometimes bears sessile dichasia of 2-3 flowers in a

terminal spike ; by the development of branches from the lower leaf-
in other cases
axils it becomes a branched panicle. The single flowers have five stamens, and
are only slightly zygomorphic the three posterior stamens have hairy filaments,
;

and are further distinguished from the two anterior stamens by the transverse

KM;. 74'J. .YiV(>f in/in ti'lun-irm (A nut. si/c). I'Msoxiir*.

of their anthers. Diyitalis, Foxglove (Figs. 747, 748), has an obliquely

rainpanulate corolla and four stamens. The flowers hang from one side of the
obliquely ascending raceme. Scrophularia has a two-lipped corolla with a very
short lower lip. j^The corolla in Linaria and Antirrhinum is spurred. Veronica
and Gratiola have only two fertile stamens.
(b) JRhinantheae.Tbia group includes a number of closely related genera which
have adopted a moiv or less completely parasitic mode of life. The most com-
666 BOTANY

pletely parasitic form is Lathraea (cf. p. 211), the species of which have no trace of
chlorophyll ;
L. squamaria, the Toothwort, is parasitic on the roots of the Hazel.
Many (e.g. Tozzia, Bartsia, Euphrasia, Odontites, Pedicular is, Melampyrum,
Aledorolophus) are semiparasitic. Although they possess green leaves they attach
themselves by means of haustoria to the roots of other plants, from which they
obtain nutrient materials. Further details regarding the mode of life and develop-
ment of these forms, and the seasonal dimorphism they exhibit, will be found in
ss
the literature cited ( ).

OFFICIAL. Diyitalis purpurea yields DIGITALIS FOLIA.

Family 3. Orobanchaceae. Root-parasites, without chlorophyll. Flower as

V\<: 743. Nicotiana taltncum. a, Flower ; b, corolla, cut open and spread out flat ; c, ovary ;

rf and e, young fruit, (a, d, c, nat. size ; e, d, x 2.)

in the Scrophulariaceae, but with a unilocular ovary. Several British species of

Orobanchc, parasitic on various host plants (Fig. 749).
Family 4. Bignoniaceae. Woody plants, in many cases climbers. Stamens 4,
corolla two-lipped. Seeds winged. Catalpa bignonioides, Tecoma radicans, orna-
mental plants from X. America.
Family 5. Gesneriaceae. Tropical herbs with a unilocular ovary which is some-
times superior, in other cases inferior, and 4 or 2 stamens. Frequently cultivated
on account of the beautiful colour of their flowers, e.g. Gloxinia, Aeschynanthus,
Achimenes. Ramondia pyrenaica is one of the few European representatives.
w
Streptocarpus ( ) polyanthus only develops one of its cotyledons, which persists
and attains a large size ;
the inflorescences arise at the base of the cotyledon.
 PHANEROGAM I A 667

Fio. 744. Hyoscyamus niger. Flowering shoot and fruit (4 nat. size). OFFICIAL and Pomoxoi's.

FIG. 745.Scrophulariaceae. Floral diagrams. Fio. 746. Verbuvum thapsiforme. a, Flower ;

A, Verbascum ; B, Gratida. (After EICHLER.) b, calyx and style (nat. size).

668 BOTANY

Family ti. Lentibulariaceae. Marsh- or water-plants with a spurred corolla

and only two M ),
(anterior) stamens. They capture and digest insects. Utricularia (

aquatic plants provided with bladder-like traps in the tropics also terrestrial and
;

epiphytic forms. Pinguicula, with a rosette of radical leaves, the margins of which
are inrolled (cf. Fig. 47 and p. 237).
Family 7. Herbs with decussately arranged leaves and quad-
Acanthaceae.
rangular stems. The and spring open when mature.
fruits are bilocular capsules
Numerous species occur as undergrowth in tropical forests. Species of Acanthus
occur in the Mediterranean region. Their large, pinnately-lobed leaves, the lobes
terminating in spines, figure in Grecian art on the capitals of pillars, etc.

FIG. 747.JH<jit(ilis i>urpurea. a, Flower ; b, corolla cut open and spread out c, calyx ;ui<l pistil
; ;

d, fruit after dehiscence ; e, transverse section of fruit (nat. size). OFFICIAL.

Family 8. Plantaginaceae. Herbs with a basal rosette of leaves and long-

stalked spikes of crowded flowers. Flowers tetramerous, actinomorphic. Corolla
membranous. Ovary bilocular. Style long and papillate.
GENERA. The native species of Planlago ( 66 ), the Plantain, are markedly proto-
gynous (cf. Fig. 241, p. 310) the young heads or spikes show the pale-coloured
;

stigmas projecting before the corolla has opened (Fig. 750). Transitional forms
from anemophily to entomophily are found. Litorella lacustris is an aquatic
plant with a spike of three flowers two female flowers are situated at the base,
;

while the male flower has a long stalk.

Order 7. Rubiinae

Plants of diverse habit with opposite leaves. In contrast to the

preceding order the flowers have a sympetalous corolla associated
 PHANEROGAMIA G69

with an inferior ovary. Flowers tetramerous or pentamerous.

Fio. M8. Digitalis intrpurea (.} nat. size). OFFICIAL and Poisosous.
 670 BOTANY TAUT II

I-' ic.. 749. Orobanche minor, parasitic on Trijiiliinn < Via. TOn. -
mnlin
/'/,>/,/,(,/,, ( nat. sixc'). SingU; (lower
(J nat. si/i-). Single flower (enlarged). and frnlt (enlarged),
 PHANEROGAMIA 671

Ovary bi- or tri-locular. Stamens epipetalous. Calyx very incon-

spicuous.
Family 1. Rubiaeeae. Herbs, shrubs, or trees, with simple,
decussate leaves and stipules. Flowers actinomorphic. Ovary
bilocular.

I 'K;. 701. Cinchona suceirulru (n.it. size). OFFICIAL. (After SCHUMANN and ARTHUR MEYEK.)

SUB-FAMILIES AND IMPORTANT GENEKA. (a) Cinchoneae, woody plants with

scaly stipules and numerous seeds in each cavity of the ovary. Cinchona is a
genus from the S. American Andes, now cultivated in the mountains of nearly all
tropical colonies
751, 752).
(Figs. Stipules deciduous. Flowers in terminal
panicles ; tubular, with an expanded terminal portion fringed at the
corolla

margin. Fruit, a septicidal capsule, the valves of which, after dehiscence has
taken place, are still held together above by the small calyx. Seeds with a broad,
672 BOTANY

membranous wing. Ourouparia ( 57 ) is a genus of climbing plants found in the

Malayan Archipelago the hook-like organs specialised for climbing are the stalks
;

of reduced inflorescences, which are sensitive to contact. The flowers are associated
together in spherical inflorescences resembling those of the commonly grown orna-
mental shrub Cephalanthus.
(b) Coffeae. Stipules scaly. Each loculus of the ovary with a single seed.
Coffea, the Coffee plant, is a shrub C. arabica (Fig. 753) and C.
; liber ica are

FIG. 752. A, Flower.

CiTichona succirubra. B, Corolla split oi-n. (.', Ovary in longitudinal
section. D, Fruit. E, seed. (D, nat. size,the others enlarged.) After A. MEYER and
SCHUMANN. OFFICIAL.

important economic plants, originally derived from Africa, and now cultivated
throughout the tropics. The pentamerous, snow-white flowers, which have a
similar scent to that of Orange blossom, and the bright red fruits contrast with
the shining, dark-green foliage, and make a coffee plantation attractive both in
appearance and in scent. The pericarp becomes differentiated into a succulent
exocarp and a stony endocarp. The minute embryo is embedded in the horny
endosperm with which the seed is filled ; very thin. The remark-
the seed-coat is

able myrmecophilous plants (cf. Fig. 207, p. 237) of the Malayan Archipelago,
58
Myrmecodia ( ) and Hydnophytum, also belong to this group the former has ;

spiny roots. Uragoga Ipecacuanha, a small herbaceous plant or undershrub, is a

native of 'Brazil, especially of the province of Matogrosso. The stipules have an
 PHANEROGAMIA 673

incised margin, and the inflorescence is a terminal head of flowers surrounded by

a number of bracts (Fig. 754). The fruits are small berries. The plant is

cultivated in suitable localities in valleys on the south side of the Himalayas.

(c) Stcllatae. This is the only subdivision of the family which is represented
in Europe ; Galium, fiubia, Aspcrula (Woodruff). These genera are characterised
by the resemblance of the stipules to the leaves (Fig. 755) usually a whorl of ;

six members is borne at each node, but sometimes the number is increased, and in

!'";. "53. Coffea arabica(% nat. size). Single flower, fruit, seed enclosed in endocarp, and freed
from it (about nat. size).

Rubia it is reduced to four by the union of the stipules in pairs. The leaves are
easily distinguished from the stipules by the presence of buds in their axils.
Flowers usually tetramerous a single seed in each loculus of the ovary. Fruit
;

a schizocarp splitting into nut-like, partial fruits. Fruits often provided with
hooks.
OFFICIAL. Cinchona succirubra yields CINCHOXAK RUBKAE CORTEX. QUININE
is obtained from this and other species of Cinchona. Uragoga (Psychotria)
//>< cm-iianha yields IPECACUANHA. CATECHU is obtained from Ourouparia
( Uncaria) gambir.
2 x
674 BOTANY

Family 2. Caprifoliaeeae. Woody plants, usually without

stipules. Flowers sometimes zygomorphic. Ovary usually trilocular.
Fruit a berry or drupe.
SUB-FAMILIES AND IMPORTANT GENERA. (a) Sambuceae, with actinomorphic

FIG. 754. Uragoga Ipecacuanha (J nat. size). Infrutescence by the side. OFFICIAL.

flowers and drupes. Viburnum, leaves simple, fruit one-seeded. Sambucus, Elder,
with imparipinnate leaves, glandular stipules, and three stones in each fruit.
S. nigra (Fig. 756) is a common European shrub. The flat-topped inflorescences
are panicles consisting of decussately arranged main branches which ultimately
bear the flowers in small cymes.
(b) Lmiicereae, flowers zygomorphic, fruit a berry. The Honeysuckle (Lonicera
 PHANEROGAMIA 675

Diervilla ( Weigeiia) is a favourite

flowering shrub.
OFFICIAL. Sambucus nigra yields
SAMBUCI FLORES.

Family 3. Valerianaeeae.
Herbs or small shrubs, with de-
cussately arranged leaves and
asymmetrical flowers. Calyx
only developed with the fruit as
a pappus. Corolla pentamerous,
spurred. Stamens 4, 3, or 1.
Carpels 3, but only one loculus of
the ovary is fertile, and contains
a single, pendulous ovule (Figs.
757, 758).

GENEKA. Valeriana, Valerian. V.

officinalis is a herbaceous plant widely
distributed throughout Europe. It
has a short, thick, chambered rhizome,
which often sends out subterranean
offsets. Leaves imparipinnate. In-
florescence a panicle. Spur short. The
hairy calyx developed on the single-
seeded fruit assists in its dispersal by
the wind. Valcrianella, small, incon-
spicuous plants. V. olitoria, Lamb's
Lettuce. Centranthus ruber from the
Mediterranean region has a long spur
to the flower ;
it is commonly culti-
vated in gardens.
OFFICIAL. Valeriana officinalis
yields VALERIANAE IIHIZOMA.

Order 8. Campanulinae

Herbs, with simple, entire,

alternate, exstipulate leaves.
Floral formula usually 5, C(5), K
A 5, G
(2-3); in other cases
there are 5 carpels. A well-

developed calyx, and stamens

inserted on the floral axis with Fio. 755. Asperula odoruta (jj nat. size). Single
their anthers coherent are char- flower and flower in longitudinal section (en-
larged).
acteristic of the order.

Family 1. Campanulaceae. Herbs with milky juice flowers actiuomorphic ;

;

ovary as a rule trilocular. Fruit a capsule (Figs. 759, 760).

676 BOTANY PART II

FIG. 756. Sambucus nigra. Flowering shoot and infrtitesoence (A nat. size).
Single flower (enlarged).OFFICIAL.

FIG. 757. Valeriana. Floral Fiii. "58. Valeriana officinalis. tt, Flower (x h. fruit
ts) ;

diagram. (x 4). OFFICIAL.

 PHANEROGAMIA 677

BRITISH GEXKUA ( 59 ). Campanula, numerous species with showy flowers.

Phytcumn, with spike-like inflorescences. The petals separate at the base, remaining
o

V\<;. 7">!t. Floral diagram of Campanula FIG. 760. Campanula rotundifolia. a, Flower b, the ;

medium. (After EICHLEB.) same cut through longitudinally. (Nat. size.)

united at the tip until the pollen, which is shed in the bud, is swept out by the
hairs on the style after this the lobes of the
;

stigma, which until now have been closed, ex-

pand. Jasione resembles the Compositae in
having its flowers in heads and its anthers
united to form a tube.
Family 2. Lobeliaceae. Herbs with milky
juice, closely related to the Campanulaceae.
Flowers zygomorphic carpels 2. The median
;

sepal is anterior and comes below a deep in-

cision in the corolla. The normal position is
assumed by torsion of the whole flower through
180 or inversion of the flower these methods ;

also occur in the Orchidaceae (Fig. 761). In

Britain Lobelia Dortmanna, an aquatic plant
of northern regions which reaches its southern
limit in the north of Germany.
OFFICIAL. Lobelia inflata from N. America
(Fig. 762) yields LOBELIA.

Fie. 701. Floral diagram of Lobelia fulgens. (After Fi<:. 7i'>2. Lobelia inflata nat. size).
(
Kli FII.DR.) OFFICIAL.

60
Family 3. Cucurbitaceae ( ). Herbs, without milky juice, climbing by
678 BOTANY PART II

means of tendrils the cordate or palmately lobed leaves bear coarse hairs.
;

Flowers diclinous moncEcious or less commonly dioBcious.

; Calyx and corolla
adherent below. Anthers united in pairs or all coherent, CO-shaped. Ovary
trilocular. Fruit a berry, with a firm rind and six large, succulent, parietal
placentas.
IMPORTANT GENERA. Cucumis sativus, the Cucumber, and C. Melo, the Melon,
are commonly cultivated. Tendrils un-
branched the tendril corresponds to a bract
;

and arises nearer to the apex of the shoot

than the leaf nearly opposite to which it
stands. The female flowers are solitary,
the male are grouped in small inflorescences,
in the axils of the leaves. As has been
shown by NOLL, fertilisation is not a neces-

sary preliminary to the development of the

fruit of the Cucumber. Cucurbita pepo, the
Pumpkin, has branched tendrils, which re-
present axillary shoots of the simple tend-
rils. Each branch of the tendril corresponds
to a leaf of the shoot, and may sometimes
subtend an axillary bud. Both male and
female flowers are solitary in the leaf-axils.
Bryonia, the Bryony, is British (Fig. 764) ;

it is poisonous and has thick, swollen,

fleshy roots, long unbranched tendrils, and
lla
axillary inflorescences ( ). Citrullus Colo-
cynthis is a perennial plant inhabiting the
Asiatic and African deserts north of the

FK;. 764. Bryonia dioica. A, Y\o\\> in..: i

branch (reduced); B, female, C, male

FIG. 763.
A'c6aijim(Cucurbitaceae). Diagram of flower (nat. size); D, androecium (niaxiii-
a male (A) and of a female flower
(). (After fled); E, fruits; t\ fruit in section.
EICHLER.) Ponoiroot.

equator. Leaves deeply three lobed and Tendrils simple or

pinnately divided.
forked male and female flowers The fruit
;
solitary in the axils of the leaves.
isa dry berry (Fig. 765).
OFFICIAL. Citrullus colocynthis ELATKKU M
yields COLOCYNTHIDIS PULPA.
from Ecballium elaterium.
SECT. II PHANEROGAMIA 679

FIG. ~65.Cilnillus culocynthis (J nat. size). 1, Shoot with male and female flowers. 2, Apex of a
shoot with a male flower-bud and tendrils. 3, Male flower with corolla spread out. k, Female

flower cut through longitudinally. 5, Young fruit cut transversely. OFFICIAL.

Order 9. Aggregatae
For the most part herbs with the flowers in heads, surrounded by
a common involucre. In relation to this the calyx of the individual
680 BOTANY PART II

flowersis reduced. Stamens epipetalous. Ovary inferior, unilocular

with a single ovule. Fruit indehiscent.

Family 1. Dipsacaceae. Herbs with opposite leaves. Flower with an epiralyx

a c I*

FIG. 766. Snccisa pratensis. a, Flower with cpiralyx ;

FIG. 767. Compositae. Floral
b, the same after removal of epicalyx c, fruit in ;
diagram (Canluus).
longitudinal section ; /, ovary ; hk, epicalyx.

which persists on the fruit. Tetramerous or pentamerous. Stamens 4, anthers

free. Ovule pendulous, anatropous. Seed with endosperm.
IMPORTANT GENERA. Dipsacus, the Teazel, lias recurved hooks on the involucral
and floral bracts. The capitula of D. fullonum are employed in the carding of
woollen cloth. Corolla with four lobes. Sitccisa (Fig. 766) has a four-lobed corolla ;

bracts are present on the common recep-

tacle. Scabiosa has similar bracts but
has pentamerous flowers the marginal ;

\ x flowers of the head are larger and

x-..
^^^^^ *^!lf/
dorsiventral. It is cultivated as an
ornamental plant. Knautia has tetra-
merous flowers no floral bracts.
;

C1
Family 2. Compositae ( ).
For the most part herbs of very
various habit some tropical ;

forms are shrubs or trees.

Flowers actinomorphic or zygo-
a C T& morphic with no epicalyx.
|.-], ; . Arnica montana.
TilS. a, Ray-flower; l>. Stamens five J
anthers intrOl'SC,
dis-flower; e, the latter cut through longitudin-
cohering by their Cuticles tO
ally. (After BERO and SCHMIDT, magnified.)
, , , , .
,
,

form a tube which is closed

below by the unexpanded stigma. The pollen is shed into the tube
formed by the anthers and is swept out by the brush-like hairs of
the style as the latter elongates. The style is bifid above. Ovule
erect, anatropous (Fig. 770). Seed exalbuminous. The fruits are
achenes, often bearing at the upper end a crown of hairs, the pappus.
This corresponds to the calyx and aids in the dispersion of the
 PHANEROGAMIA 681

fruit by the wind (Figs. 767-773). The replacement of reserve

starch by inulin is characteristic of the Compositae.
The individual flowers are either radially symmetrical with a five-lobed corolla
(Fig. 768 b, c) or they are two-lipped as in the South American Mutisieae, the

FIG. 769. Longitudinal section of capitulum a, of Lappa major with floral bracts 6, of Matricaria
;

Chairwmilla without floral bracts. (After BERG and SCHMIDT, magnified.)

upper lip having two teeth, the lower three. By suppression of the upper lip
flowers with a single lip are derived such flowers exhibit three teeth at the tip (Fig.
;

768 a). The

ligulate flowers (e.g. of Taraxacum, Fig. 775) are similar in general
appearance to the latter the corolla is here deeply split on one side and its
;

margin bears five teeth. In addition to those Compositae which have only
ligulate or only tubular florets in
the head, there are many which have
tubular florets (disc-florets) in the
centre, surrounded by one -
lipped
florets (ray
-
769 b).
florets) (Fig.
These usually differ from one another
in sex as well as in colour the disc- ;

florets are hermaphrodite, the ray-

florets purely female. The flower-
heads are thus heterogamous (Matri-
caria, Arnica). Lastly, the marginal
florets may be completely sterile
(Centaurea Cyanus) and serve only
to render the capitulum conspicuous
to insects ^ I0- Arnica montana. a, Receptacle of capi-
^"^-
,
,.
.,. tulum removal of fruit; h, fruit in longi-
after
Important sub-families and genera. tudina , 8ectfon> the pappl]8 on y partly shown
,

1. Tiibulifiorae, the heads contain only (After BERO and SCHMIDT, magnified.)
tubular florets :
(a) Cynareae. The en-
larged axis of the inflorescence or common receptacle has setaceous floral bracts ;

the involucral leaves form several series and are

prickly or have membranous
margins. The
flowers are either all hermaphrodite or the marginal florets
are sterile. Style swollen below the stigmas into a cushion-like ring. Fruits
with a pappus. Carduus (Plumeless Thistle), pappus of simple, hair-like bristles.
Cirsium, with feathery pappus. Echinops, with single-flowered capitula associated
682 BOTANY

in numbers. Lappa (Burdock), involucral bracts with recurved, hook-like tips

(Fig. 769 a). Cynara Scolymus (Artichoke). Cnicus benedictus (Fig. 774), capit-
ula solitary, terminal, surrounded by foliage leaves. In-
volucral bracts with a large, sometimes pinnate, terminal
spine and a felt of hairs. Marginal florets sterile.

Pappus double. Centaurea with dry, scaly involucral

bracts and large, sterile marginal florets. On the irrit-
ability of the stamens cf. Fig. 238 and p. 296.
(b) Eupatorieae. Flowers hermaphrodite. Receptacle
without floral bracts. Involucral leaves herbaceous.
Eupatorium cannabinum (British). Aggcratum mexi-
canum, commonly cultivated.
2. Afutisieae occur in tropical America.
Labiatiflorae.
3. Cichorieae with only zygomorphic,
Liguliflorae.
ligulate, florets with a five-toothed corolla in the flowcr-
head. All the flowers are hermaphrodite. Laticiferous
vessels in all organs of the plant. Taraxacum ojficinale
(Dandelion, Fig. 775) is a common plant throughout
the northern hemisphere. It has a long, carrot-shaped

tap-root, a rosette of coarsely toothed leaves, and terminal

inflorescences, borne singly on hollow stalks (apogamy,
cf. p. 518). Fruits with an elongated beak, carrying
up the pappus as a stalked, umbrella-shaped crown of
hairs (Fig. 773). Lactuca sativa, Lettuce. L. virosa,
a herbaceous plant of considerable height with a branched
stem, sessile sagittate leaves and numerous, small, few-
flowered inflorescences grouped in panicles. Fruits like
Fn;. 771. Androecimii of
Carditus crispiis (x 10).
those of Taraxacum. L. Scariola C52 ), Compass plant,
(After BAILLON.) has leaves which take a vertical position. Cichorium
Intybus (Chicory), C. endivia, Endive, with heads of blue
flowers ; pappus of short erect scales (Fig. 772 C). Tragopoyon and Scorzoncra have
a feathery pappus ;
Sc. Crepis has a soft, flexible, hairy pappus of
hispanica.
M
brownish colour. Sonchus, pappus, of several series of bristles. Hieracium ( ),

7?

Fio. 772. Fruits of A, Helianthvs a unit us ; Ji, Hieracium virosum ; C,

Intybus. (After BAILLON.)

a large European genus with many forms. (Apogamy and apospory, cf. p. 517.)

Pappus white, rigid, and brittle (Fig. 772 B).

4. Diver siflorae. Two distinct types of flower or flowers of distinct sexes in the
capitulum.
 PHANEROGAMIA 683

FIG. 773. Head of fruits of Taraxacum ojflcinale. The pappus is raised on a .stalk-like

prolongation of the fruit. (Xat. size.)

FKI. 774. Cnicus benedictw. (After BAILLON.)

684 BOTANY I'.VUT II

l-'ii:. 775. Taraxanim officinale (i nat. siz<-). liy th<- side is a rcrfj.tacli' still Ix-ariiii: twr) ,
if tin-
fruits ; the i>a])])iis is raist-il on a stalk-like Iwak of the fruit OFFICIAL.
 PHAXEROGAMIA 685

() Astereae, receptacle naked, without floral bracts. Involucral bracts, numer-

FIG. 777. Artemisia

Cina. (After SCHU-
MANN and ARTHUR
I i<;. n 9,Matnaana Chamomilla ( nat siae). MEYER )

ous, imbricate. Hermaphrodite tubular flowers in centre, surrounded by one-

686 BOTANY

lipped ray-florets. Pappus as a rule of bristle -like hairs. Style with straight
branches stigmas terminal. Numerous species of Aster, Solidago, and Eriyeron
;

occur in Europe, America, and Asia. Species of Aster are cultivated. Baccharis ( M ),
m
shrubby dioecious plants from America. Species of Jfaastia ( ) are cushion-shaped
plants with woolly hairs in New Zealand (Vegetable Sheep).
(b) Inuleae resemble the preceding group but have tailed anthers. Involucral
leaves frequently dry and membranous. Inula occurs in Britain ;
/. helenium

Fio. 778. Tussilago Farfara. (After BAILLON.)

is cultivated.In Gnaphalium, Antennaria O*5*), Hclichrysum (Everlasting

flowers),Leontopodium (Edelweiss), Filago, etc., the dry involucral bracts are
coloured and have flat, expanded appendages ; the flowers are all tubular but
the marginal florets are female, the disc-florets hermaphrodite. Raoulia ( 66 )
resembles Haastia in habit and in distribution (cf. Fig. 193). Odontospermum
pygmaeum (Rose of Jericho) closes its iuvolueral bracts over the fruits when dry
and expands them when moistened.
(c) Heliantheac. Receptacle with floral bracts. Involucral bracts herbaceous.
 PHANEROGAMIA 687

Disc -florets
hermaphrodite, tubular. Ray-florets one -lipped, female, or her-
maphrodite. Anthers without tails. Pappus wanting or formed of awns or
scales. Helianthus annuus (Sunflower, Fig. 772 A), If. tuberosus (Jerusalem

FIG. 779. Arnica montana (J nat. size). OFFICIAL.

Artichoke), Dahlia, are all from America and in cultivation. In Britain Bidens
herbs with opposite leaves, sometimes heterophyllous. Espeletia ( 67 ), characteristic
Paramos of Colombia and Venezuela.
trees or herbs of the Tagetes, Zinnia, orna-
mental plants from America.
688 BOTANY PART n

(d) Anthemidcae. Receptacle with or without floral bracts or hairs ;

involucnil
bracts with dry, membranous margins ;
in other respects like tin- preceding.
Achillca, Milfoil Anthemis, Anacyclus, with floral bracts.
; Anthem-is imliilix.
or with these more or less replaced by
capitula composed of disc-florets only,
irregular florets. Anacyclus qfficinarum. Afatricaria has no floral bracts.
M. Chauiomilla (Chamomile, Figs. 769 b, 776) is an annual, copiously branched
herb with a hollow, conical, common receptacle, yellow disc-florets arid white,
recurved, female ray-florets, iu the terminal capitula. Chrysanthemum, <'. */// VH.//I,
Tanacetum, flowers all tubular, marginal florets female. Artemisia has all the
florets tubular and usually the peripheral ones female (A. Abxiiitliitun. Worm-
wood) in the few-flowered capitula of A. Cina (Fig. 777) all the florets are
;

hermaphrodite.
(e) Senecioneae. Receptacle naked, involucral bracts in one or two series.
Pappus hairy, delicate, white in other respects like the preceding. Tussilayu
;

Farfara, Coltsfoot, flowers appear before the leaves the flowering stem bears
;

scaly leaves and a single capitulum (Fig. 778). Female flowers at periphery in
several series. Leaves large, cordate, thick, covered beneath with white hairs.
Petasitcs offieinalis, Butter-Bur. Senecio, plants of diverse habit, including some
trees and succulent plants of world-wide distribution.
; vulgaris has no ray-
>S'.

florets but only tubular, hermaphrodite florets. Doronicum, Cineraria are com-
monly cultivated. Arnica montana (Figs. 779, 768, 770) has a rosette of radical
leaves in two to four opposite pairs and a terminal inflorescence bearing a single

capitulum ;
from the axillary buds of the two opposite bracts one (rarely more)
lateral inflorescence develops.

(/) Calenduleae. Fruits of the capitulum of varied and irregular shapes.

Calendula.
OFFICIAL. Anacyclus Pyrethrum yields PYHETHKI HADIX. SANTOXIXV.M is
prepared from Artemisia maritima, var. Stechmanniana. Anthemis nobilis yield>
AXTHEMEDIS FLOKES. Taraxacum qfficinale, TARAXACI KADIX. Arnica montana,
AUXICAE RHIZOMA.

6
Fossil Angiosperms ( ")

The first undoubted Angiosperms appear in the Upper Cretaceous. They are
represented by numerous species which, like the recent forms, can be divided into
Monocotyledons and Dicotyledons. The most ancient forms are known only as
leaves, so that their determination is a matter of difficulty. They agree essentially
with living Angiosperms, and since they show no similarities to Gymuosperms or
Pteridophytes, do not aid in bridging over the gap between the Augiosperms and
these groups.
Of Monocotyledons examples of the Palmae are known from the Cretaceous
period onwards. The leaves of Dicotyledons from the Cretaceous are doubtfully
placed in various orders which contain the more lowly organised types (Querciflorae,
Platanaceae) some are ascribed to Eucalyptus. The Angiosperms of the Eocene
;

and the Oligocene can be determined with greater certainty even in Northern ;

Europe representatives of existing tropical and sub-tropical families occurred,

e.g. Palmae, Dracaena, Smilax among Monocotyledons, numerous Querciflorae

(esp. Quercus), Lauraceae (Cinnamonum,

etc.), Leguminosae among Dicotyledons.
As the present period approached the fossil Angiosperms throw even less
ia

light on the systematic arrangement of the group. The few genera that have
 PHANEROGAMIA

become extinct (e.g. Dryophyllum, a supposed ancestral form of the Oaks) are
wanting even in the Eocene. From the Miocene onwards the specific forms are in
part identical with those now living, and in the Quaternary strata all the remains
are of existing species. The general character of the Tertiary flora in Europe was,
however, very different from that of the present day. It had the aspect of the
flora of a much warmer region, and (as in the case of the -Gymnosperms) contained
forms which now exist only in distant regions.

2 Y
 INDEX OF LITEBATUKE

INTRODUCTION AND MORPHOLOGY

2
( ) FRITZ
!
( ) E. HAECKEL, Generelle Morphologic der Organismen, 1866, p. 52.

MULLER, Fiir Darwin, 1864. ( ) CHARLES DARWIN, On the Origin of Species by

3

( ) HUGO DE
4
Means of Natural Selection, 1859. TRIES, Die Mutationstheorie,
5
1901-1903. ( ) CARL NAGELI, Mechanisch-physiologische Theorie der Abstam-
mungslehre, 1883. JULIUS SACHS, Flora, voL Ixxxii. 1896, p. 173. ( 6 ) R. v.
WETTSTEIN, most recently in Handbuch der systematischen Botanik, 1901, p. 40.
7 "
( ) Cf. John Butler-Burke in Nature," 1905, pp. 78, 294, 492 this investigator's ;

results have been widely published in the daily press. On the other side see
especially the review by W. Roux, Wochenschrift iiber die Fortschritte und
Bewegungen auf dem Gesamtgebiete der Wissenschaft und Technik, 1906, No. 8.
8
( ) Another interpretation is given by K. GOEBEL, Organography of Plants,
1898-1901 [English translation, 1900-1905]. 9
) K. GOEBEL, loc. cit. and
(

Vergleichende Entwickelungsgeschichte der Pflanzenorgane, 1883 F. PAX, ;

10
Allgemeine Morphologic der Pflanzen, 1890. ( ) K. GOEBEL, Organography,
H K. 12
vol. i. p. 19, vol. ii. p. 35. ( ) GOEBEL, loc. cit. vol. i. p. 42. ( ) K. GOEBEL,
13
) K.
Biol. Zentralbl. vol. xxii. 1908, p. 388. ( GOEBEL, Organography, vol. ii.
p. 241. (
14
) A. W. EICHLER, Zur Entwicklungsgeschichte des Blattes, 1865.
15 16
( ) K. GOEBEL, Organography, vol. ii. p. 308. ( ) M. RACIBORSKI, Flora,
vol. Ixxxvii. 1900, p. 1. n ) v. DEINEGA, Flora, vol. Ixxxv. 1898, 18
( p. 439. ( )

Ann. du Jard. Bot. de Buitenzorg, vol. ii. 1893, p. 98. ( 19 ) K. GOEBEL, Organo-
20
) S. SCHWEN-
21
graphy, vol. ii. p. 350. t ) K. GOEBEL, Bot. Ztg. 1880, p. 753. (

DENER, Mechanische Theorie der Blattstellungen, 1878, together with numerous

C ) HANS WINKLER, Jahrb. f.
22
papers in the Stzber. d. Akad. d. Wiss. zu Berlin.
wiss. Bot. vol. xxxvi. 1901, p. 1, and vol. xxxviii. 1903, p. 501 further literature
;

in these p) WIESNER, Biol. Zentralbl. vol. xxiii. 1903, 209. M

papers, p. ( )

K. GOEBEL, Arb. d. bot. Inst. in Wiirzburg, vol. ii. 1882, pp. 357 ff., and Biol.
Zentralbl. vol. xxii. 1902, p. 389. M ) FREIDENFELT, Flora, vol. xci. 1902,
( p. 115.
26
( ) ROSTOWZEW, Flora, vol. xlviii. 1890, p. 155, and K. GOEBEL, Organography,
) A. F. W. SCHIMPER, Die epiphytische Vegetation Amerikas,
27
vol. ii. p. 226. C
28
1888. EUG. WARMING, most recently in ENGLER and PRAXTL, Nat. Pflanzen-
C )
29
familien, Teil, Abt. 2a, 1891, p. 2.
3. ( ) K. GOEBEL, Flora, vol. xcv. 1905,
31
p. 165. C* ) K. GOEBEL, Organography, vol. ii. p. 226. ( ) Cytologische Studien

aus dem Bonner botauischen Institut, Jahrb. f. wiss. Bot. vol. xxx. 1897, p. 155.
FR. MEVES, Verhandl. d. Anat. Gesell., sechzehnte Vers., Halle, 1902, p. 152.
M. KOERNICKE, Der heutige Stand der pflanzlichen Zellforschung., Ber. d.

691
692 BOTANY

deutsch. bot. Gesellsch. vol. xxi. (1903), p. 82. KOERNICKE, Centrosomen bei
den Angiospermen ? Flora, vol. xcvi. 1906, p. 501. C 32 ) Cf. on the one hand
ALFRED FISCHER, Untersuchungen liber den Bau der Cyanophyceen und Bacterien,
1897, and Die Zelle der Cyauophyceen, Bot. Zeitg. vol. Ixii. 1905, p. 51, and on
the other F. G. KOHL, iiber die Organisation uud Physiologic der Cyauophyceeuzelle
and EDGAR W. OLIVE, Beihefte zum Bot. Zentralbl. vol. xviii. 1904, p. 9. M> (

HUGO BE w
VRIES, Jahrb. f. wiss. Bot. vol. xvi. 1885, p. 465. ( ) N. GAIDUKOV,
M ) Most
Ber. d. deutsch. bot. Ges. 1906, p. 155. ( recently PAUL KKETZSCIIMAK,
Jahrb. f. wiss. Bot. vol. xxxix. 1904, p. 273. (*) FR. MEVES has given the name
of " chondromites to thicker fibrils of peculiar nature which he has observed in
"

certain cases in the cells which line the pollen-sacs. Ber. d. deutsch. Bot. Ges.
1904, p. 284. Cf. also RUDOLF BEER, Beiheft zum bot. Zentralbl. vol. xix. 1905,

p. 304, and G. TISCHLER, Jahrb. f. wiss. Bot. vol. xlii. 1906, p. 568. ( 37 ) Cf. on
this subject especially ALFRED FISCHER, Fixierung, Farbung und Bau des Proto-

plasma, 1899, and ALBERT DEGEN, Bot. Ztg. 1905, Abt. I. p. 202. t ) Cf. the most
38

recent works of GREGOIRE and his pupils as well as those of E. STRASBURGEU.

(*) BELAJEFF, Three papers on Sperm atogenesis in the Ber. d. deutsch. bot. Ges.
1897, p. 337 if.; IKENO, Jahrb. f. wiss. Bot. vol. xxxii. 1898, p. 557; HIRASE,
Journal of the College of Science Tokyo, vol. xii. 1898, p. 105 ; HERBERT J.
WEBBER, Botanical Gazette, vol. xxiii. 1897, p. 453, vol. xxiv. 1897, p. 16 and
p. 225 ; S. IKENO, Beihefte zum bot. Zentralbl. vol. xv. 1903, p. 65. K. MIYAKE,
Bot. Magaz. Tokyo, vol. xix. 1905, p. 98, and Ber. d. deutsch. bot. Gesell. 1906,
p. 78. (
40
) A.
F. W. SCHIMPER, Jahrb. f. wiss. Bot. vol. xvi. 1885, p. 1, and Bot.
41
Ztg. 1880, p. 886. ( ) L. MARCHLEWSKI, Section Chlorophyll in vol. viii. of

ROSCOE-SCHORLEMMER, Organ. Cheinie, 1901, p. 83 C. A. SCHUNK, Proceed. Roy. ;

Soc. London, vol. Ixv. 1900, p. 177, vol. Ixviii. 1901, p. 474, and vol. Ixxii. 1904,
42
p. 165 also A. TSCHIRCH, Ber. d. deutsch. bot. Gesell. 1904, p. 414.
; ( )
43
TSCHIRCH, Untersuch. lib. d. Chlorophyll, 1884. ( ) L. MARCHLEWSKI, cf. in*
41 also W. KUSTEE, Ber. d. deutsch. bot. Gesell. 1904, p. 339
;
L. MARCHLEWSKI, ;

ibid. 1906, p. 146. (**) TH. W. ENGELMANN, Bot. Ztg. 1882, p. 663 HANS ;

MOLISCH, Bot. Ztg. 1905, Abt. I. p. 131. (

45
) HANS MOLISCH, in 44. C
46
) C. A.
SCHUXK in the last edition of 41, p. 172 ; cf. MOLISCH, Ber. d. deutsch. bot.
Gesell. 1896, p. 27 ; TAMMES, Flora, vol. Ixxxvii. 1902, p. 205, and KOHL, Unters.
47
iiber d. Karotin, etc. 1902; A. TSCHIRCH, in 41, p. 417. ( ) Complete literature

in Louis GAUCHER, Etude generale sur la membrane cellulaire chez les vegetaux,

PFEFFER, Unters. aus dem bot. Inst. zu Tiibingen, vol. i. 1885, p. .''25
48
1904. ( ) ;

G. HABERLANDT, Sinnesorgane im |Pflanzenreich, 1901, p. 126. C* ) W. ROTHERT,

9

Anzeiger der Akad. d. Wis. in Krakau, 1897, p. 11. C ) G. HABERLANDT, Die

Lichtsinnesorgane der Laubbliitter, 1905. Cf. also H. R. VON GUTTENBERG, Ber.
51
d. deutsch. bot. Gesell. 1905, p. 265. ( ) Numerous researches of JULIUS WIESNER,

collected in his Anatomic und Physiologic der Pflanzen, edition ir. 1898, p. 39 if.
and p. 339. C2 ) GILSON, La cristall. de la cell, in La Cellule, vol. is. 1893, p. 397.
53
( ) L. MANGIN, Journ. de Bot. vol. vii. 1893, pp. 37, 121, 325. Complete
literature to 1904 in Louis GAUCHER in 47. More recently FRIEDRICH CZAPEK,
Biochemie d. Pflanz. 1905, vol. i. p. 506. ( w ) GILSON, Rech. chim. sur la Membr.
cell, des Champ, in La Cellule, vol. xi. 1894, p. 7.
M ) FR. CZAPEK in
( Hoppe-
Seyler's Ztschr. f. physiol. Cheinie, vol. xxvii. 1899, p. 141 Congres internal, de ;

Bot. Paris, 1900. (

M) F. C. VON FABER,
Ber. d. deutsch. bot. Gesell. 1904, p. 177.

(
87
) VAN WISSELINGH, Archives Neerland. vol. xxvi. 1892, and vol. xxxviii. 1894.
(
M ) Cf. especially Z. KAMERLING, Bot. Zentralbl. vol. Ixxii. 1897, p. >85. (*)
KLEBS, Unters. aus dem
60
bot. Inst. zu Tubingen, vol. i. 1885, p. 582. t ) Litera-
 INDEX OF LITERATURE 693

61
ture in ERNST KUSTER, Pathol. Pflanzenanat. 1903, p. 10. ( ) MOLISCH, Studien
82
iiber Milchsaft und Schleirnsaft der Pflanzen, 1901. ( ) CARL NAGELI, Die
Starkekorner, 1858. (*') A. F. W. SCHIMPER, Bot. Ztg. 1881, p. 223 ARTHUR ;

MEYER, Unters. iiber die Starkekorner, 1895 cf. also SYNIEWSKI, Bull, internat. ;

de 1'Acad. d. sc. de Cracovie, 1899, p. 245, who there criticises A. Meyer's

distinction of two physical modifications of the substance of the starch-grain.
(
M H.
) FISCHER, Beitr. z. Biol. d. Pfl. vol. viii. 1898, p. 53. (
M HENRY KRAEMER,
)
6
Bot. Gazette, 1902, p. 341. C* ) L. MAQUENNE and Eire. Roux, Compt. rend.
67
1905, vol. cxl. p. 1303, and 1906, vol. cxlii. p. 95. ( ) 0. BUTSCHLI, Verh. d.
68
naturhist.-med. Ver. at Heidelberg, 1903, vol. xvii. p. 451. ( ) A. TSCHIRCH and

II. KRITZLER, Ber. d. deutsch. pharm. Gesell. 1900, vol. x. p. 214. ( W. 69

)

PFEFFER, Jahrb. f. wiss. Bot. vol. viii. 1872, p. 472. ( 70 ) A. ZIMMERMANN, Beitr.
71
z. Morph. u. Phys. der Pflanzenzelle, vol. i. 1891, p. 113. ( ) L. KNY, Ber. d.
72
deutsch. bot. Gesell. 1887, p. 387. ( ) M. RACIBORSKI, Anzeiger, Akad. Wiss.
73
Krakau, 1893, ( p.
) A. TSCHIRCH, Ber. d. deutsch. Bot. Gesell. 1904,
259.

p. 419. (
74
BUTSCHLI in 67, pp. 436 and 506.
) Cf. 0. (
75
) LEO EKRERA,
des and 7S
L'epiplasme Ascomycetes, 1882, other papers. ) G. BERTHOLD, Studien (

iiber Protoplasmamechanik, 1886, p. 24. (") A. TSCHIRCH, Die Harze und die
Harzbehalter, 1900. (
78
R. CHODAT and A. BACH, Archives, des sciences phys.
)

et math, de Geneve, 1904, etc.; then W. PALLADIN, Ber. d. deutsch. bot. Gesell.

1905, p. 240, 1906, p. 97 most recently T. KNASNOSSELSKY, in Ber. d. deutsch.

;

79
bot. Gesell. 1906, p. 134. C ) M. RACIBORSKI, Ber. d. deutsch. bot. Gesell. 1898,
pp. 52, 119, and Flora, 1898, p. 362. WINOGRADSKY, Bot. Ztg. 1887, p. 493,
and Beitrage zur Morph. und Phys. der Bakterien, 1888. ( 81 HANS MOLISCH,
O
)

) On the botanical side the numerous works of

82
Bot. Ztg. 1905, Abt. I. p. 161. (

E. STRASBURGER, M. TREUB, L. GUIGXARP, WL. BELAJEFF, J. BRETLANP

FARMER, NEMEC, V. GREGOIRE, BERGHS, ROSENBERG, ALLEN, MIYAKE, J. B.
OVERTON and others. (
83
) TH. BOVERI, Ergebnisse iiber die Koustitution der
chromatischen Substanz des Zellkerns, 1904 ;
V. GRKGOIRE and A. WYGAERTS,
84
Beihefte z. bot. Zentralbl. vol. xiv. 1903, p. 18. ( ) E. STRASBURGER, ALLEN,
MIYAKE, J. B. OVERTON, Jahrb. 1906, p. 1 ; the various f. wiss. Bot. vol. xlii.

investigations on the nature of the reduction of chromosomes should be compared

in the works of the authors mentioned in 82. (
M ) HARPER, SWINGLE, STRAS-
BURGER, Jahrb. f. wiss. Bot. vol. xxx. 1897 ; MOTTIER, Ber. d. deutsch. bot.
Gesell. 1898, p. 124 ; MOORE, ibid. p. 266.(*) R. A. HARPER, Jahrb. f. wiss.
Bot. vol. xxx. 1897, p. 249. (
8T
) On the botanical side numerous works especially
of G. THURET, N. PRINGSHEIM, E. STRASBURGER, L. GUIGNARD, NAWASCHIN,
and others. ( K ) Especially the works of WL. BELAJKFF, most recently in Ber. d.
89
deutsch. bot. Gesell. 1897, p. 337 ft'. ( )
The works cited under 39 by IKENO,
90
HIRASE, J. WEBBER, and K. MIYAKE. C ) E. STRASBURGER, Jahrb. f. wiss. Bot.
91
vol. xli. 1905, p. 88. ( ) JUEL, Botan. Zentralbl. vol. Ixxiv. 1898, p. 369. Abh.
(* ) Sw. MURBECK, Lunds
2
d. Kl. Schwed. Akad. d. Wiss. vol. xxxiii. 1900, No. 5.
Univ. Arskrift, vol. xxxvi. Afd. II. No. 7 Kongl. Fysiogr. ; Siillsk. Handlingar,
vol. xi. No. 7, 1901 E. STRASBURGER in 90. ; (
93
) JAMES BERTRAM OVERTON,
94
Bot. Gazette, vol. xxxiii. 1902, p. 363. ( ) C. RAUNKIAER, Archiv fiir Botanik,
95
vol.ii. N. 4, p. 7 0. JUEL, Botan. Tidsskr. vol. xxv. 1903, p. 109.
; C ) C. H.

OsTENFELi) and C. RAUNKIAER, Bot. Tidsskr. vol. xxv. 1903, p. 409, and Ber. d.
deutsch. bot. Gesell. 1904, p. 376 ;
0. ROSENBERG, ibid. 1906, p. 157. () AL.
BRAUN, Abhandl. Berl. Akad. 1856, p. 337
d. A. DE BARY, Bot. Ztg. 1857, ;

97
p. 379.C ) SHAW, Bot. Gazette, vol. xxiv. 1897, p. 114; ALEX. NATHANSOHN,
Ber. d. deutsch. bot. Gesell. 1900, p. 99. C*) Of the copious literature there need
 694 BOTANY

only be mentioned AHTHUK MEYEK, in Bot. Ztg. 1896, Abt. I. p. 187, and Ber. d.
deutsch. bot. Gesell. 1897, continued in recent years of
p. 166 ;
W. GARDINER,
the Proc. of the Roy. Soc. London, and Proc. of the Cambridge Phil. Soc.; E.
STKAsijt'RGER, Jahrb. f. wiss. Bot. vol. xxxvi. 1901, p. 493, the whole literature
is given in this paper ;
ARTHUR W. HILL, Phil. Trans. Roy. Soc. London, vol.
cxciv. 1901, p. 83, and Annals of Botany, vol. xv. 1901, p. 575. (") A. DE BARY,
Vgl. Anat. d. Vegetationsorgane, 1877 [Translated. Comparative Anatomy of
the Phanerogams and Ferns] G. HABERLAXPT, Physiol. PHanzenanat. 3rd edition, ;

1896 ; H. SOLEKEPER, Syst. Anat. d. Dicotyledonen, 1899. ( 10 ) HENRI DEVAUX,

101
Mem. de la Soc. des Sc. phys. et nat. de Bordeaux, vol. iii. 1903, p. 89. ( )

H. KHOEMER, Bibl. Bot. Heft 59, 1903, p. 20. ( ) G. HABERLANDT in m 50. (

m )

E. STRASBURGER, Jahrb. f. SCHWENDENER,

wiss. Bot. vol. v. 1866, p. 297 ;
S.
104
Monatsber. d. Berl. Akad. d. Wiss. 1881, p. 883, etc. ( ) G. HABERLANDT,
Phys. Pflanzenanat. 3rd edition, 1896, p. 430, Bot. Unters. SCHWEXDEXEU,
1<w
dargebracht, 1899, p. 104. ( ) HABKRLAXDT, Sinnesorgane ini Pflanzenreich,
106
1901. ( A. NESTLER, Ber. d. deutsch. bot. Ges. 1900, pp. 189, 327 PKTKR*,
) ;

107
Stzber. d. Niederrh. Gesell. Bonn, Med. Sekt. 1900, p. 33. ( ) In 101, p. 26.
loe I0st
( ) Cf. N. WILLE, Beitrage zur physiol. Anat. der Laminariaceen, 1897. ( )

A. G. TAXSLKY and EDITH CHICK, Annals of Botany, vol. xv. 1901, p. 13.
uo
( )

in 104, p. 308. U1 F. C. vox FABEK, Ber. d. deutsch. bot. Gesell. 1904, p. 296.
( )
ns ALFRED
( ) FISCHKR, Studien iiber die Siebrohren der Dikotylenblatter, 1885;
E. iiber den Ban und die Verrichtung der Leitungsbahnen in den
STRASBURGER,
Pflanzen, 1891, pp. 98, 297. ( ) PH. VAN TIEGHEM, Traite de Botanique, edition
m
2, 1891, p. 737 STRASBURGER, the work cited under 112 J. C. SCHOUTE, Die
Stelartheorie, 1902. (
;

m
) Against the Stelar Theory cf. especially SOLMS-LAUBACH,
;

Bot. Ztg. 1903, Abt. II. pp. 37, 147. ns

( ) HABEHLAXDT, Jahrb. f. wiss. Bot. vol.
xxxviii. 1902, p. 450. (
116
) in 56, p. 181. ( ) G. HABERLAXDT, in 104, p. 244.
m
(
118
) C. SAUVAGEAU, M. vox MINDEX, P. WEIXROWSKI, cf. in G. HABERLANDT,
n9
Physiol. PHanzenanatomie, 3rd edition, 1904, pp. 437, 475. ( ) In NAGELI,
Beitrage zur wiss. Bot. Heft I. 1858, and especially in the work of DE BARY,
iao
cited in 99, p. 243 ff. ( ) Literature in A. GRAVIS, Mem. publ. par. 1'Acad. roy.

de Belgique, vol. Ivii. 1898 R. STERCKX, Arch. Inst. bdt. univ. Liege, vol. ii.
;

) W. EICHLER, Sitzber.
121 122
1900. ( ) J. C. SCHOUTE, Flora,
(
vol. xcii. 1903, p. 32.
d. Berl. Akad.
Wiss. vol. xxviii. 1886,
d. M. BARSICKOW, Verb. d. phys. p. 501.
med. Ges. zu Wiirzburg, 1901, vol. xxxv.
p. 213. ( ) Especially the works of DE
m
BARY cited under 99, of E. STRASBURGER under 112, of HABERLANDT under 99,
and the works of BURGERSTEIX in the publications of the Wien. Akad. ( 124 ) J. C.
SCHOUTE, Verhandl. d. Koninkl. Akad. van Wetensch. Amsterdam, Tweede Sectie,
Deel IX. No. 4, 1902. SCHOUTE terms a cambium active on both sides dipleuric,
to one side only monopleuric ;
a cambium witli a single initial layer is spoken of
as initial cambium in contrast to stratified cambium proceeding from several
12S
contributory layers. H. MAYR, Zeitschr. f. Forst- u. Jagdwesen, vol. xxv.
( )
126
1893, pp. 313, etc. ( ) ALFRED FISCHER, Jahrb. f. wiss. Bot. vol. xxii. 1890,
p. 73. E. STRASBURGER in 112, p. 883 ff. (
127
L. KXY, Ber. d. deutsch. bot. )

Gesell. 1890, p. 176, and E. STRASBURGER under 112. 128

( ) Especially PH. VAX
TIEGHEM, Traite de Botanique, 2nd edition, 1891, p. 719. Also I. FREIDENFELT,
Flora, vol. xci. 1902, p. 115, and M. BUSGEN, Flora, vol. xcv. 1905, p. 58. (
12S
)

Especially H. SCHEXCK, Beitriige zur Biologic und Anatomic der Lianen, 1892.
13
) A. B. FRANK, Die Krankheiten der Pflanzen, 2nd edition, 1895, vol. i.
( p. 81 ;

KUSTER in 143. ( 131 ) J. C. SCHOUTE under 124. ( 132 ) A. B. FRANK, Bot. Ztg.
1864, p. 186 G. KRAUS, Abh. d. naturf. Ges. zu Halle, vol. xvi. 1885,
;
p. 365 ;
 INDEX OF LITERATURE 695-

E. STRASBURGER ;
R. MEISSXER, Bot. Ztg. Abt. I. 1894, p. 55, and
in 112, p. 107
Especially DK BART in 99, p. 560 DAMM, Beihefte z. bot,
133
1901, p. 25. ( ) ;

1S4
Zentralbl. vol. xi. 1901, p. 222. ( ) HANNIG, Bot. Ztg. Abt. I. 1898, p. 24.
13a
( ) STAHL, Bot. Ztg. 1873, p. 561 H. DEVAUX, Ann. des sc. nat. Bot. ser.
;

1900, vol. xii. p. 1 ; literature in HABERLANDT in 99, p. 419.

136
8, ( ) H. v.
MOHL, Bot. Ztg. 1860, pp. 1, 132, 273 ; v. BRETFELD, Jahrb. f. wiss. Bot. vol. xii.
1879-81, p. 133 ; VAN TIEGHEM, Traite de Bot., Edition 2, 1891, vol. i. p. 865 ;
STABY, Flora, 1886, p. 158 ; TISSON, Mem. de la Soc. Linn, de Normandie,
vol. xx. 1900. (
m
) Cf. especially J. WIESNER, Ber. d. deutsch. bot. Gesell. 1906,
138
p. 32. ( ) Literature in the work of KUSTER cited under 143. (
139
) Cf. B.

NEMEC, Studien iiber die Regeneration, 1905. More recently \V. FIGDOR, Ber. d.
deutsch. bot. Gesell. 1906, p. 13. (
14
) HENRI DEVAUX, Actes de la Soc. Linn, de
141
Bordeaux, ser. 6, vol. viii. 1903, p. 98. ( ) OTTO APPEL, Ber. d. deutsch. bot.
142
Gesell. 1906, p. 118. ( ) A. TISON, Bull, de la Soc. Linneenne de Normandie,
se'r. 5, vol. viii. 1904, p. 176. (
143
)
HANS WINKLER, Ber. d. deutsch. bot. Gesell.
1902, p. 81 K. GOEBEL, Biol. Zentralbl. vol. xxii. 1902, p. 385 ; Flora, vol. xcii.
;

1903, p. 132 ; ERNST KUSTER, Pathologische Pflanzenanatomie, 1903, p. 8 ; Fu.

HILUEBRAND, Ber. d. deutsch. bot. Gesell. 1906, p. 39. ( 144 ) H. LEITGEB, Unter-
suchungen iiber die Lebermoose, 6. Heft, 1881. (
145
) AUGUSTS PYRAME UK
146
CANDOLLE, Theorie elementaire de Botanique. ( ) DE WILDEMAN, Memoire
la E.
147
couronnes, etc. publics par 1'Acad. de science de Belgique, 1898, vol. liii. ( )

H. LEITGEB in 144, 3. Heft 1877; this includes the literature. ( 148 ) W. HOF-
MEISTER, Die Lehre von der Pflanzenzelle, 1867, p. 135 includes the older ;

literature later in numerous special papers by LEITGEB

;
on Equisetu.ni cf. also ;

149
E. STRASBURGER, Bot. Praktikum, edition 4, p. 312, includes literature. ( ) J.

SACHS, Arbeiten des Bot. Inst. in Wiirzburg, 1878 and 1879, pp. 46, 185. vol. ii.
lso
( ) JOHANNES HANSTEIN, Festschr. d. niederrh. Gesell. f. Natur- und Heilkunde,
151
1868, p. 109. ( ) L. ERRERA, Bull, de la soc. Beige de Microscopie, vol. xiii.
p. 12, 1886 and in 60. Vers. Naturforscher u. Arzte zu Wiesbaden, Biol. Zentralbl.
;

1887-88, p. 728 ; BERTHOLU, Studien iiber Protoplasmamechanik, 1886, p. 219 ;

cf. also DE WILDEMAN in 146. (
152
) C. NAGELI and LEITGEB, Beitr. z. wiss. Bot.

von C. NAGELI, Heft, 1868, p. 73.

4. ( ) ROSTOWZEW,
153
Flora, vol. xlviii. 1890,
134
p. 155. ( ) Especially E. v. JANCZEWSKI, Ann. des sc. nat. Bot. ser. 5, vol. xx.
1874, pp. 162, 208, and PH. VAN TIEGHEM, Traite de Bot. 2nd edition, 1891,
15B
p. 694, where the literature is given. ( ) K. GOEBEL, Organography of Plants,
vol. ii. p. 242. (
156
) LUDWIG KOCH, Jahrb. f. wiss. Bot. vol. xxv. 1893, p. 380.
Especially PH. VAN TIEGHEM in 154, p. 700, where the literature is given.
157
( )
158 159
) The most important
( ) K. GOEBEL, Organography of Plants, vol. i. p. 141. (

literature will be found in the works of A. WEISMANN, TH. BOVERI, E. STRAS-

16
BURGER. Especially 0. PENZIG, Pflanzen-Teratologie, 1890, where the
( )

literature will be found K. GOEBEL, Organography of Plants, vol. i. p. 177.

;

ERNST KUSTER in 143 WERNER MAGNUS, Ber. d. deutsch. bot. Gesell. 1903,
;

p. 129. ERNST KUSTER terms abnormal development of the cell contents or of

the thickening of the wall of the cells metaplasy, excessive division of cells
hyperplasy, and arrest in the process of development hypoplasy.

PHYSIOLOGY
For more advanced study the new edition of PFEFFER'S Physiology of Plants,
as well as JOST'S Lectures on Plant-Physiology, both of which are translated into
English, and the lists of literature contained in them may be consulted.
 96 BOTANY

On
separate aspects of Physiology further information will be found in AD.
MAYER, Agrikulturchemie, 5th edition, Heidelberg, 1901, 1902; HABERLAMT.
Physiologische Pflauzenanatomie, 2nd edition, Leipzig, 1896 SCHIMPER, Plant- ;

Geography, English edition, Oxford, 1903 HOMER, Physikal. Cheraie d. zelle u.

;

d. Gewebe, Leipzig, 1902 CZAPEK, Biochemie d. Pflanzen, Jena, 1905.

;

!
( ) MEZ, Flora, xciv. 1905, includes literature.

The law of VAN 'T HOFF, that the rapidity of a chemical reaction is about doubled
by raising the temperature 10", has also been verified for some physiological pheno-
mena. In the organism, however, there are usually anti-reactions which conceal
the primary dependence on the temperature, and give rise to the well-known
cardinal points with an optimum which is soon reached. Literature in revit-w l>y
JOST, Biol. Zentralbl. xxvi. 1906.
According to BECQUEREL (Compt. rend. 140, 1905) dry seeds can withstand
without injury temperatures of - 190.
On the geographical distribution of plants see SCHIMPEK, Plant-Geography,
Eng. ed. 1903. DRUDE, Handb. der Pfl.-Geogr. 1890. GRISEBACH, Veget. d. Erde,
1872. ENGLER, Versuch einer Entwicklungsgesch. d. Pflanzemvelt, Leipzig, 1879
and 1882. SOLMS-LAUBACH, Leitende Gesichtspunkte d. allg. Pfl.-Geogr. Leipzig,
1905. IHNE, Phanolog. Karte des Friihlingseinzugs in Mittelenropa. ;
Peterni.
Mitt. 1905, Heft 5. HOCK, Ankbmmlinge in d. Pflanzenwelt ;
Beihefte Bot.
Zentralbl. II. xviii. 1904.
(-) ANDREWS, Jahrb. f. wiss. Bot. xxxviii. 1902. MOTTIER, Annals of Bot.
xiii. 1899. MIEHE, Flora, vol. Ixxxviii. 1901. FRUH, Abbildung d. vorherr-
schenden Winde durcli die Pflanzemvelt, Zurich, 1901-2. (
:!

)PFEFFER, Osmotische
Untersuchungen, 1877. DE VRIES, Jahrb. f. wiss. Bot., vol. xiv. ;
ibid. vol. xvi.
Botan. Ztg. vol. xlvi. 1888, and vol. xlvii. 1889. (
4
) SI-HWEXDEXER, Das mech.
Prinzip im anatom. Ban der Monokotylen, 1879.
5
( ) HABERLANDT, Physiolog.
6
Pflanzenanatomie, 2nd edition, 1896, p. 134 fF. TSCHIRCH, Flora, xciv. 1905. ( )
7
AMBROXN, Jahrb. fur wiss. Bot. vol. xii. 1879. ( ) F. Sen WAR/, Phys.
Unters. lib.
Dickenwachst. u. Holzqual. von Piuus silv., Berlin, 1899. HARTIG, Holzunter-
suchungeu, Berlin, 1901. WIEDERSHEIM, ibid, xxxviii. 1903. 0. MELVILLE
BALL, Jahrb. f. wiss. Bot. xxxix. 1903. VOCHTING, Nachr. Kgl. Ges. d. Wissensch.
Gottingen M. N. Kl. 1902, Heft 5. URSPRUNG, Ber. dtsch. Bot. Ges. 1901.
SONNTAG, Jahrb. fur wiss. Bot. xxxix. 1903. HARTIG, Holzuntersuch., Berlin,
1901, p. 53. WILDT, Inaug.-Diss., Bonn, 1906. ( 8 ) NOLL, Thiel's Landw. Jahrb.
vol. xxix. 1900, p. 361. 9
( ) NAGELI, Denkschr., schweiz. Naturf. -Ges. vol. xxxiii.
1893. MOORE and KELLERMANN, U.S. Dep. of Agricult. 190">. MA-AYASU KANDA,
Journ. Coll. of Sc. Tokyo, vol. xix. (Review, SORAUER, Ztschr. f. Pfl.-Krankh.
10
15.) ( ) WINOGRADSKY, Ann. de 1'Inst. Pasteur, 1890, 1891. Arch. d. sc. biol.
Inst. imp. d. Med. exper. a St. -Petersb. 1892. Zentralbl. f. Bakteriol. 1896.
STUTZER, Mitteil. d. Landw. Instituts d. Univ., Breslau, 1898. BEHREXS, Arbt.
d. Bakt. im Boden, etc. Arb. deutsch. Landw. Gesell. 1901, Heft 64. LOHXIS,
Zentralbl. f. Bakt. II. xiii. 1904. BEXECKE and KEUTXER, Ber. dtsch. Bot. Ges.
xxi. 1903. KEUTNER, Wiss. Meeresuntersuch. vol. viii. Kiel, 1904. (") Literature
in A. FISCHER, Vorles. liber Bakterien, 2nd edition, Jena, 1903, and in HILTNKK in
:

LAFARS, Handb. d. technischen Mykologie, 2nd edition, vol. iii. BENECKK and
KKUTXER, Ber. dtsch. Bot. Ges. 1903, xxi. REIXKK, Ber. dtsch. Bot. Ges. xxi.
1903, and xxii. 1904. (
12
) SAIDA, Ber. dtsch. Bot. Ges. 1901, xx. J. KUHN,
Fiihlings Landw. Ztg. 1901, p. 1. WAUMBOLDT, Unters. iiber d. Biol. stickstoff-
bind. Bakterien. Inaug.-Diss., Gottingen, 1905. HASELHOFF and BREDEMAXN,
13
Landwirtschaftliche Jahrbiicher, xxxv. 1906. ( ) BENECKE, Bot. Ztg. 1903, i.
 INDEX OF LITERATURE 697

79. v. PORTHKIM, Sitzber. Wien. Akad. vol. ex. I.

(
u
) v. D. CRONE, Sitzber.
]>.

Niederrh. Ges. f. Natur- u. Heilkunde, Bonn, 1902, and Inaug.-Diss., Bonn, 1904.
(
13
) RADLKOFER, Ber. dtsch. Bot. Ges. xxii. 1904. ROTHERT (Alumin.), Bot. Ztg. i.

xciv. 1906. GOSSL (Mangan.), Beiheft Bot. Zentralbl. xviii. I. 1905. ( 16 ) WIEGMANX
and POLSTORFF, tJber die anorgan. Bestandteile d. Pflanzen, 1842. Cf. v. LIPPMAXX,
in Chemikerzeitung, 1894. (
17
)
E. WOLFF, Aschenanalysen von land- und forst-
18
wirtsch. Produkteu 1871 and 1880. ( ) N. WILLE, Festschrift f. Scliwendener,
19
1899, p. 321. ( ) PFEFFER, Landw. Jahrb. 1876. Osmotische Untersuchungen,
1877. Zur Kenntnis d. Plasmahaut u. d. Vakuolen, Abh. math. -phys. Klasse
Kgl. Sachs. Ges. d. Wiss. 1890. Uber Aufnahme u. Ausgabe ungeloster Kiirpcr :

20
ibid, and Pfl.-Phys., 2nd edition, vol. i. chap. iv. t ) NATHAXSOHX,
Ber. dtsch.
21
Bot. Ges. 1901, p. 509. PULLST, Jahrb. f. wiss. Bot. xxxvii. 1902. ( ) G.
SCHRO-
DER, Uber die Austrocknungsfahigkeit d. Pflanzen, Inaug.-Diss., Leipzig, 1886.
AL. BRAUX, Betrachtungen iiber die Verjiingung i. d. Natur, 1850, p. 213. KURZ-
WELLY, Jahrb. f. wiss. Bot. xxxviii. 1903. FRITZ MULLER, Kosmos, vol. xiii.
1883, and Prometheus, vol. ix. Heft 1. (*>) THIEL, Landw. Zentralbl. 1870.
NOBBE, Versuchsstat. 1875. POND, U.S. Fish, commiss. Rep. for 1903, Washington,
1905. BUSGEX, Flora, xcv. 1905. Ergbd. RACIBORSKI, Bull. Acad. d. sc.,
Cracovie, 1905. KUXZE, Jahrb. f. wiss. Bot. xlii. 1906. O23 ) HALES, Ess. of veg.
statics, 1727. G. KRAUS, Flora, 1882, and Forschung. a. d. Gebiet d. Agrikult.-
Phys. 1887. CHAMBERLAIX, Rech. s. 1. seve ascend., Bull. du. Lab. d. Bot. ge"n.
de 1'Univ. de Geneve, vol. ii. 1897. PITRA, Jahrb. f. wiss. Bot. 1877. C24 ) FIGDOR,
25
Sitzber. Wien. Akad. d. Wiss. cvii. 1898. MOLISCH, Bot. Ztg. i. 1902. C )

STRASBURGER, Leitungsbahneu, Jena, 1891, p. 537. v. HOHXEL, tfber d. negat.

Druck der Gefassluft, Inaug.-Diss. Strassb. 1876, and Jahrb. wiss. Bot. 1879.
NOLL, Sitzber. niederrh. Ges., Bonn, 1897. STRASBURGER (Luftverdiinnung), I.e.

p. 712. DIXON and JOLY, Ann. of Bot. 1895. Report of a Discussion on the ascent
of 26
water, British Assoc. Liverpool, Sept. 1896. C ) ASKEXASY, Verhandl.
naturhist.-med. Verein Heidelberg, 1895 and 1896. STEIXBRIXCK, Ber. dtsch. Bot.
Ges. 1902. Dixox, Scient. Proc. Roy. Soc. Dublin, vol. x. 1903. COPELAXD,
Bot. Gaz. xxiv. 1902. SACHS, Arb. Botan. Institut Wiirzburg, vol. ii. and Ges.
Abhandl. i.
p. 23. KAMERLIXG, Bot. Zentralbl. 1898. URSPRUXG, Beihefte z.
Bot. Zentralblatt, xviii. 1904, and xix. 1906, Jahrb. f. wiss. Bot. xlii. 1906. STEIN-
BRIXCK, Flora, xciv. 1905, and Jahrb. f. wiss. Bot. xlii. 1906. ( ) ASKEXASY, A'erh.
-27

nat. med. Ver., Heidelberg, 1896. STRASBURGER, Leitungsbahnen. ( 28 ) SCHWEN-

UEXEU, Monatsber. Kgl. Akad. d. Wiss., Berlin, Juli 1881, and Sitzber. d. Akad.
<1. Wiss., Berlin, 1889. LEITGEB, Mittlg. d. Bot. Inst., Graz, 1886. STAHL, Bot.
Ztg. 1894. KOHL, Bot. Beiblatt d. Leopoldina, 1895. FR. DAUWIX, Phil.
Transact. Roy. Soc., London, vol. cxc. ser. B. p. 531. COPELAXD, Ann. of Bot.
Ixii. 1902. BURCKRSTEIX, Die Transpiration der Pflanzen, Jena, 1904, includes
29
literature. ( ) v. HOHNEL, Mittlg. a. d. forstl. Versuchswesen Osterreichs, 1879,
and Forsch. a. Gebiet
Agrikult.-Phys. 1881.
d. HABERLAXDT, Wiss.-prakt.
d.
Unters. a. d. Gebiete des Pflanzenbaues, 1877. BUSGEX, Ban u. Lebeii der Wald-
) Cf. STAHL and DAUWIX in 27, BUSCALIOXI and POLACCI,
:to
biiunie, 1897, p. 168. (
31
Atti del Istit. hot. dell' Univ. de Pa via, vii. 1902. ( ) TREUB, Ann. Jard. Bot.,

Buiterizorg, 1889. KRAUS, Flora, vol. Ixxxi. 1895. KOORDERS, Ann. Jard. Bot.,
Buitenzorg, 1897. MOLISCH, Ber. dtsch. Bot. Ges. xxi. 1903. ( 32 ) GOEBEL, Flora,
Ixxvii. 1897. PERCY GROOM, Ann. of Bot. 1897. HABERLAXDT, Jahrb. f. wiss. Bot.
34 35
1897, p. 27. ( ) NOLL, Flora, 1893. ( ) SCHIMPER, Die epiphytische Veget.
Amerikas, 1888, and Indo-malayische Strandflora, 1891. SCHIMPER, Pflanzen-
geographie, Jena, 1898. GOEBEL, Pflanzenbiolog. Schilderungen I. KIHL.MAXX,
698 BOTANY

x ) Cf. PFEFFKK,
Pflanzenbiolog. Untersuchungen in Russisch-Lappland, 1891. (

Pfl.-Physiol. 2nd edition, vol. i. 60. BEYERIXCK, Progr. de la soc. holl. des
sciences a Harlem, 1904. The results of KOHL, Bi-r. dtscli. Bot. Ges. 1897, Heft 2,
37
that the ratio is 5 : 4, is due to an error in calculation. ( ) EXGELMAXN, Bot. Ztg.
1884, 1887, 1888, Verb. Akad. Amst. 1894. REIXKE, Bot. Ztg. 1886. EXGEI..M ANN.
Arch. f. Anat. u. Physiol. 1902. GAIDUKOW, Abh. Berl. Akad. d. Wiss., Anhang
1902. PAXTAXELLI, Jahrli. f. wiss. Bot. xxxix. 1903. REINKE, Lehrb. f. wiss. Bot.
x. MOLISCH, Bot. Ztg. Ixiii. 1905, i. Abt. TSWETT, Bot. Ztg. Ixiii. 1905.
1876.
TSWETT, Ber. dtsch. bot. Ges. xxiv. 1906. KOHL, Ber. dtsch. bot. Ges. xxiv. 1906.
38
STAHL, Naturw. Wochenschr. 1905. No. 45. t ) DETLEFSEN, Arb. Bot. Inst.,
Wiirzburg, 1888, vol. iii. HOR. BROWX, Address to the Chem. Sect. Brit. Assoc.
for advance of Science, 1900. KEIXKE (Ergriinen), Sitzber. Preuss. Akad., Berlin,
1893. LOB, Landwirtsch. Jahrb. xxxv. 1906, Heft 4.
EXGELMAXX and GAIDUKOW found that Oscillaria sancla assumed a red
colour in green light, a green colour in red light, a blue-green colour in yellow
light, and a brownish-yellow colour in blue light. They term this property com-
plementary chromatic adaptation, and see in it a confirmation of the action of
pigments as sensibilisators.
39
( ) The criticism in an English review that the intermediate products are not

evident from this equation, rests on a misunderstanding of the character of chemical

equations. According to SCHLOESIXG, Compt. rend. vol. cxxxi. more is liberated

than C02 decomposed.

40
( ) KREVSLER, Landw. Jahrb. 1885. GODLEWSKI, Arb. Bot. Inst. Wiirzburg,
vol. i. H. BROWN, cf. under No. 32 BUOWN and ESCOMBE, FARMER and CHANDLER,
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MATTHAEI, Phil. Trans. Roy. Soc. Lond., cxcvii. 1904. BLACKMAN, Ann. of
Bot. xix. 1905. BLACKMAX and MATTHAEI, Proc. Roy. Soc. Lond., Ixxvi. 1905.
DEMOUSSY, Compt. rend, cxxxix. 1904.
According to a recent result CO can be utilised when the percentage present is

considerable (?).

(
41
) H. BROWN
and ESCOMBE, Static Diffusion of Gases and Liquids in relation
to the assimilation of Carbon and translocation in Plants, Phil. Trans. Roy. Soc.
42
ser. B. 193, 223, 1900. ( ) F. HOFMEISTER,
Die chemische Organisation der Zellc.
Braunschweig, F. Vieweg u. Sohn, 1901. (Naturw. Rundschau xvi. 1901.) G.
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xxvii.

According to STEPHAN 1 mgr. NaCl requires about 1 year to penetrate from a

10% solution for a distance of 1 metre into water, cane sugar would require 24
years, and albumen fully 14 years. The greater rapidity of osmotic movements in
the organism therefore presupposes acceleratory influences.
(*) W. OSTWALD. Uber Katalyse. Naturw. Rundschau, xvi. Jahrg. 1901.
HOFMEISTER, ibid. p. 581 ff. G. BREDIG, I.e. CZAPEK, Antifermente, Ber.
dtsch. Bot. Ges. xxi. 1903. (*>) A. FISCHER, Jahrb. f. wiss. Bot. xxii. 1890.
MfJLLEU-THVRGAU, Landw. Jahrb. xi. 1882.
According to NIKELEWSKI, Beihefte Bot. Zentralbl. xix. 1905, the transforma-
tions of starch, sugar, and oil in trees not only depend on external influences, but
in large part on the internal periodicity of the vital processes.
(*) WEEVERS (Glucosides), Jahrb. f. wiss. Bot. xxxix. 1903. BEYERINCK, On
the formation of Indigo etc. Verb. Kon. Akad. v. Wetenschap. te Amsterdam,

31, Sept. 1899. On Indigo-Fermentation, ibid. 31, Miirz 1900. MARCHLEWSKI,

 INDEX OF LITERATURE 699

Bull. Acad. d. Sc. d. Crocovie, 1902. W. BUSSE, Mitt. a. d. Labor, d. Kais.

47
Gesdhts.-Amts, 1900. ( ) TREUB, (Pangium) Ann. Jard. Bot., Buitenzorg, 1895.
GUIGNARD, Bull. d. Stances Soc. nat. d'Agricult. de France, 1906. Journ. of the
Board of Agricult. 1906. MALLEVRE, Journ. de la Soc. centrale d'Agricult. de
Belgique, 1906. Mitt. d. deutsch. Landw. Gesellsch. 1906, No. 21. KNIEP
1

(Latex), Flora, xciv. 1905. Further PFEFFER, Pfl.-Phys..2. Ann ., vol. i. 78-91.
48
( ) SOLMS-LAUBACH, Bot. Ztg. 1874. Also in ENGLER-PRANTL, Rafflesiaceae,
49
1889, iii. 1. POKLO (Neottia), Flora, xcv. 1905, Ergbd. ( ) L. KOCH, Jahrb. f. wiss.
Bot. 1888, and Ber. dtsch. Bot. Ges. 1887. HEINRICHER, Jahrb. f. wiss. Bot.
1897, 1898, 1901. v. WETTSTEIN, ibid, and Osterr. Bot. Ztschr. 1897. JOHOW,
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vol. xxxi. E. LAURENT, Compt. rend. t. cxxxiii. 1901. HEINRICHER, Jahrb. f.

wiss. Bot. xxxiv. 1901 ;

ibid. xxxvii. 1902. C
50
) PERCY GROOM, Ann. of Bot. ix. 1905,
Linn. Soc. Journ. xxxi. JANSE, Ann. Jard. Bot., Buitenzorg, 1896. FRANK,
Lehrbuch, vol. i.
p. 259. KAMIENSKI, Bot. Ztg. 1881. PFEFFER, Landw. Jahrb.
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xviii. p. 127. E. STAHL, Jahrb. f. wiss. Bot. 1900, vol. xxxiv. p. 539. W.
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Bot. Garden, vol. i. No. 5, 1900, p. 419. MOLLER, Zeitschr. fur Forst- und
Jagdwesen, 1902. MOLLER, Ber. deutsch. hot. Ges. xxiv. 1906. SHIBATA, Jahrb. f.
wiss. Bot. xxxvii. 1902. HILTNER, Naturw. Ztschr. f. Land- u. Forstwirtsch. i. 1903.
51
( ) HELLRIEGEL and WILFARTH, Untersuch. liber die Stickstoffnahrung der
Gramineen u. Leguminosen, 1888. Other literature most recently collected by
HILTNER in LAFARS, Handb. d. Techn. Mykologie, 2nd edition, 3rd vol. (2. Liefg.).
2
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f.
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53
1900, Heft 45. ( ) SACHS, Bot. Ztg. 1853. DE BARY, Die Erscheinung der
Symbiose, Strassburg, 1879. SCHWENDENER, Nagelis Beitriige z. wiss. Bot. 1861,
1862, 1868, and Flora, 1872.
Uber Algentypen der Flechtenkonidien, 1869.
BORNET, Rech. sur les gonidies des lich., Ann. sc. nat. 5 e ser. tome xvii. STAHL,
Beitr. zur Entwickl.-Gesch. d. Flechten, 1877 and 1878. REESS, Monatsber.
Berlin. Akad. der Wissensch. 1871. STRASBUKGER, Deutsch. Rundschau, 1891.
ARTARI, Bull. d. sc. nat. de Moscou, 1899, No. 1. Cf. also the literature for
M 55
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SCHWENDENER, 1899. RACIBORSKI, Flora, 1900, p. 87. WARMING, Videnssk.
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RETTIG, Beihefte Bot. Zentralbl. xvii. 1904. ULE, ENGLERS, Jahrb. xxxvii. 1905,
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' '

bilder." 4 Reihe, Heft 1, 1906, and Naturw. Wochenschr. N. F. v. 1906.

(
M ) CLAUTRIAU, Mem. publ. par 1'acad. roy. de Belgique, t. lix. 1900. (
57
) AD.

MAYER, Compt. rend. 1900. i. p. 1389. GRIFFON, Compt. rend. 1900, vol. cxxx.

PURIEWITSCH, Physiol. Unters. lib. Pflanzenatmung, Jahrb. f. wiss. Bot. 1900,

vol. xxxv. p. 573.
INGENHAUSS already held correct views on the distribution of the respiratory
process. (WIESNER, JAN INGENHOUSS, Wien, 1905.)
(
M ) AD. MAYER, Poggend. Ann. vol. cxlii. 1871. WORTMANN, Arb. Bot. Inst.,
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700 BOTANY

1885. E. GODLEWSKI sen. and POLZEXH-.SX, Bull. Krak. Lit. Akad., April 1, 1901,
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123. (
M WIKOGRADSKY,
) Bot. Ztg. 1887, 1888. MOLISCH, Die Pflanze in ihrer
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Eisen, 1892.
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Weinbau u. Weinhandcl, 1902, and WORTMANX, Die wi*sensch. Grundlagen der
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61
iiber Bakterien, 2nd edition, Jena, 1903. ) REINKE, Wissenschaftl.
Meeresunters.
(

Neue Folge, vol. iii. Heft 2, p. 39. MOLISCH, Bot. Ztg., i. Abt. 1903, Heft, 1.
MOLISCH, Leuchtende Pflanzen, Jena, 1904, and Sitzber. k. Akad. Wiss. Wien, M. N.
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According to the investigations of TAPPEINER and others, fluorescent substances
in the presence of light are oxygen-activators they are thus able to initiate pro-
;

found chemical processes. (Index of literature in SCHROEDER, Bot. Ztg. 1905,

ii. Abt.)
NOLL, Arb. Bot. Inst. Wiirzburg. iii. 1888. MOLISCH, Sitzber. Wien. Akad.
52
der Wiss. ex. 1901. C ) STAHL, Ber. dtsch. Bot. Ges. 1885. K. ROSEXVIXGE,
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The formative "
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thus be reckoned among phenomena of movement. (NoLL, Biol. Zentralbl. xxiii.

1903.)
93
i ) MULLER-THURGAU, Landw. Jahrb. d. Schweiz, 1898, vol. xii. FRITZ
MCLLER, Kosmos, 6. Science, 1899, N. S. vol. x.
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NOLL (Parthenokarpie d. Gurke), Sitzber. Niederrh. Ges. f. Natur- u. Heilkunde,
Bonn, 1902.
SOLACOLU, G. R. Acad. sc.,' Paris, 1900, cxli., describes parthenocarpic fruits also
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Lilium, Lunaria, and Rhododendron.
TSCHERMAK, Ber. dtsch. Bot. Ges. xx. 1902. Regeneration: GOBEL, Biol.
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w
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im Pflanzenreich, 1878. VOCHTING, Bot. Ztg., i. Abt. Ixiv. 1906. SACHS, Arb. Bot.
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1906. MIEHE, Ber. dtsch. Bot. Ges. xxiii. 1905. (**) HARTING, Liunaea, 1847.

SACHS, Jahrb. f. wiss. Bot. 1860, and Arb. Bot. Inst. Wiirzburg, vol. i. 1874.
STREHL, Unters. Uber das Langenwachstum, etc., 1874. ASKENASY, Yerh.
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w ) NAGELI, Starkekorner, 1858. BUTSCHLI, Uber d. Ban quellbarer
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d. Wiss. Gottingen, 1896.
etc. PFEFFER, Phys. of Plants,
67
vol. i. ASKENASY, Verh. nat.-med. Ver. Heidelberg, 1878. PFITZER,
12. ( )

ibid. 1882. JANSE, Maandbl. v. Natuunvetensch. 1887. F. BEXECKE, Ber. dstch.

Bot. Ges. 1893. G. KRAUS, Ann. Jard. Bot. Buitenzorg, vol. xii. p. 196. A.
 INDEX OF LITERATURE 701

68
HOLLER, Schimper's Bot. Mitt, aus den Tropen, vii. 1895. t ) POPOVICI, Bot.
Zentralbl. 1900, vol. Ixxxi. pp. 33, 87. MIYAKE (Taraxacum), Beihefte Bot.
B9 of Plants, vol. ii. ch. vi.
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70
xiii. 1. PEDERSEN, Arb. Bot. Inst. Wiirzburg, 1874. v. LIEBENBERG, Bot.
( )

Zentralbl. 1884. PAMMER, Osterr.-ungar. Ztschr. f. Zuckerind. u. Landw. 1892.

SCHINDLER, Die Lehre vom Pflanzenbau. Wien, C. Fromme, 1896. Gen. Part,
p. 79. (
71
) STAHL, Jen.
Zeitschr. f. Naturw. xvi. 1883. VOCHTING, Jahrb. f. wiss.
Bot. xxv. 1893. GOBEL, Biolog. Zentralbl. xxiv. 1904, and Sitzber. Bayr. Akad. d.
Wiss. xxvi. 1896. Flora, xcv. 1905, Ergbd. NORDHAUSEN, Ber. dtsch. Bot. Ges. xxi.
1903. KORNICKE, Ber. dtsch. Bot. Ges. xxii. 1904, and xxiii. 1905. ( 72 ) KARSTEN,
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TO
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i. Abt. 1901. W. JOHANNSEN, Das A'therverfahren beim Fruhtreiben, etc., Jena,
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74
Ber. dtsch. Bot. Gesellsch. 1901, p. 76. ( ) STRASBURGER, Deutsche Rundschau,

1898, p. 402, 1899, p. 90. Figure of Taxodium mexicanum by TULE in Prometheus

75
xiv. 1903, p. 648. ( ) HOFMEISTER, Die Lehre v. d. Pflanzenzelle, Leipzig, 1867.
ENGELMANN, Bot. Ztg. 1881, 1883, 1886, 1887. PFEFFER, Untersuch. Bot. Inst.

Tubingen, vol. ii. ROTHERT, Flora, 1901, vol. Ixxxviii. p. 406. v. DUNGERN,
Ztschr. f.
allg. Physiol. i. 1. BULLER, Ann. of Bot. 1900. Experiments with
drops of mercury S. BERNSTEIN, PFLI'GER'S Archiv, vol. Ixxx.
: SHIBATA
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76
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N. F. xvii. 1897. RITTER, Flora, 1899, vol. Ixxxvi. p. 326. JOSING, Jahrb. f. wiss.
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77
C ) ARTHUR,
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SENN, Vortrag, Ixxxvii. Jahrvers. Schweiz. naturf. Ges. Winterthur, 1904. KUSTER,
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78
SCHROTER, Flora, cxv. 1905, Ergbd. C ) STEINBRINCK in Ber. dtsch. Bot. Ges.
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79
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80
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1899, p. 567. NMEC, Die Reizleitung uud die reizleit, Gewebe, Jena, Gust.
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Bot. xxxvii. 1902, and xxxviii. 1903. NOLL, Ber. dtsch. Bot. Ges. 1902. MIEHE, Jahrb.
702 BOTANY

t'. wiss. Bot. xxxvii. 1902. MASSAKT, Mem. cour. Acad. Belg., Bruxelles, 1902.
DARWIN, Statolith Theory, Proc. Roy. Soc. Ixxi. 1903. H. SCHROEDER, Beihefte
Bot. Zentralbl. xvi. 1904. TISCHLER, Flora, xciv. 1905. PICCARD, Jahrb. f. wiss.
Bot. si. 1904. HABERLANDT, Verb. Ges. dtsch. Naturf. u. Arzte. Allgem. Tl.
1904. HABERLANDT, Lichtsinnesorgane der Laubblatter, Leipzig, 1905. Statoliths
Jahrb. f. wiss. Bot. xlii. 1906. Conduction of Stimuli :
FITTING, in Ergebnisse der
Physiologic by ASHER and SPIRO (includes list of literature).
The tonic significance of the growing point for perception and the ensuing re-
action has been termed by MIEHE (Jahrb. f. wiss. Bot. xxxvii. 1902) tonic prevalence.
81
( ) NOLL, Flora, 1893. STEYER, Reizkriimm. bei Phycomyces. Inaug. -Diss., ,

MOLISCH, Sitzber. Wien. Akad., M.-N. Kl. cxi. 1.

ffi
Leipzig, 1902. ( ) STAHL,

Uber sog. Kompasspflanzen, Jen. Ztschr. f. Naturw. 1881. ARTHUR, Bull.

Purdue Univ. 1894. ( M ) OLTMANNS, Flora, 1892, 1897. ELFVINO, Finska Yet.-
Soc. Forhaudl. xliii. 1901. O. RICHTER, Sitzber. K. Akad. d. Wiss. Wien., M.-N.,
Kl. vol. cv. 1906. (
M ) KNIGHT in Ostwald's Klassikern, No. 62, 1895. NOLL,
Het. Ind., Leipzig, 1892. CZAPEK, Jahrb. f. wiss. Bot. 1898, vol. xxxii. p. 224.
NOLL. ibid. 1900, vol. xxxiv. p. 465. NEMEC, Ber. dtsch. Bot. Ges. 1900, p. 241,
and Jahrb. f. wiss. Bot. 1901, p. 2. HABERLANDT, Ber. dtsch. Bot. Ges. 1900,
M ) BARANETZKY, Flora, 1901, Ergbd. Ixxxix. NOLL, Arb. Bot.
p. 261. ( p. 138.
Inst., Wiirzburg, vol. iii. ix. and xiii. SCHWENDENER and KRABBE, Abh. Kgl.
Pr. Akad. d. Wiss., Berlin, 1892. NOLL, Flora, 1892. MEISSNER, Bot. Zentralbl.
1894. FITTING, Jahrb. f. wiss. Bot. xli. 1905. LUXBURG, ibid. CZAPEK, ibid. xlii.
1906. C
86
) BARANETZKY, Mem. Acad. imp. St. Petersbourg, ii. Ser. T. xxxi. 1883.
SCHWENDENER, Monatsber. Berlin. Akad. Dez. 1881. WORTMANN, Bot. Ztg.
1886. SACHS, Lectures, No. 38. H. SCHENCK, Beitr. z. Biol. der Lianen, 1892.
NOLL, Heterog. Induktion, 1892, p. 46. KOLKWITZ, Ber. dtsch. Bot. Ges. 1895.
87
( ) SACHS,
I.e. Lect. 38. NOLL, Heterog. Ind. S. 48. NOLL, Sitzber. Niederrh.
88
Ges. f. Nat.- u. Heilk., 8. Juli 1901. ( ) HARTIG, Holzuntersuclmngen. Altes
u. Neues, Berlin, 1901. Seedlings of Cucurbitaceae NOLL, Landw. Jahrb. xxx.
:

Ezgbd. 1901. Load-curvature WIESNER, Sitsber. d. K. Akad. d. Wiss. Wien,

:

M.-N. Klasse, vol. cxi. 1902. (*) WORTMANN, Bot. Ztg. 1881. M. MIYOSHI,
Bot. Ztg. 1894, and Flora, 1894. MOLISCH, Sitzber. Wien. Akad. 1893. STAHL,
Bot. Zeitg, 1880. WORTMANN, Bot. Ztg. 1883, 1885. J. AF KLERKER, Ofversigt
Kongl. Vetensk. Acad. Forh. 1891. BRUNCHORST, Bot. Zentralbl. 1885, vol.
xxiii. HEGLER, Verb. Ges. Deutsch. Naturf. u. A'rzte, 1891. NEWCOMBE, Science
1901. JUEL, Jahrb. f. wiss. Bot. 1900, vol. xxxiv. p. 507. BERG, Lunds Univ.
Arsskrift 35, No. 6. SAMMET, Jahrb. f. wiss. Bot. xli. 1905. Galvanotropism :

GASSNER, Bot. Ztg. 1906, i. Abt. EWART and BAYLISS, Proc. Roy. Soc. Ixxvii.
1905. SCHELLENBERG, Flora, xcvi. 1906. (*>) SACHS, Arb. Bot. Inst. Wiirzburg,
vol. ii. 1882 (1879). NOLL, Het. Ind., Leipzig, 1892, pp. 12, 35. CZAPEK,
Jahrb. f. wiss. Bot. 1898, vol. xxxii. p. 188. NOLL, Jahrb. f. wiss. Bot. 1900,
vol. xxxiv. p. 459. BARANETZKY, Flora, 1901. Erganz.-Bd. 89. ( 91 ) NOLL, Flora,
92
1893. ( ) SCHWENDENER, Monatsber. Berl. Akad. Dec. 1881. 0. Miiller, Ranken
d. Cucurbitac. Inaug. Diss., Breslau, 1886. (COHN'S Beitr., vol. iv. Heft 2.)
(93) PKEFFER, Unters. Bot. Inst. Tubingen, vol. i. HABERLANDT, Sinnesorgane,
Leipzig, 1901. FITTING, Jahrb. f. wiss. Bot. xxxviii. 1903. ( M ) NORDHAUSEN, Jahrb.
95
f. wiss. Bot. 1899, voL xxxiv. p. 236. PEIRCE, Ann. of Bot. 1894. ( ) PFEFFEB,
Pflanzenphys. Unters., Leipzig, 1871, and Periodische Bewegungen, etc. 1875.
OLTMANNS, Bot. Ztg. 1895. JOST, Jahrb. f. wiss. Bot. 1898. WIEDERSHEIM,
Jahrb. f. wiss. Bot. xl. 1904. BURGERSTEIN, Jahrber. Erzherzog Rainer-
Gymnasium, Wien, 1902. FARMER, New Phytologist, London, vol. L
 INDEX OF LITERATURE 703

Chemonasty WACHTER, Ber. deutsch. hot. Ges. xxiii. 1905.

: (**) See under
9o. JOST and SCHWENDENER, Sitzber. K. Preuss. Akad. d. Wiss. 1897 and 1898.
OT
MOLISCH, Ber. dtsch. Bot. Ges. xxii. 1904. ) STAHL, Bot. Ztg. 1897, p. 71. (

A. FISCHER, Bot. Ztg. 1890. NOLL, Het. Ind. 1892, p. 9. (") SACHS, Flora, 1863.
KABSCH, Bot. Ztg. 1861, 1862. DARWIN, Bewegungsvermbgen, 1881, p. 271.
PFEFFER, Pfl.-Physiol. 1881, vol. ii. 59. SACHS, Lectures, No. 34.
w ) HABER- (

LANDT, Das reizleitende Gewebesystem der Sinnpflanze, Leipzig, 1890. MAC-

DOUGAL, Botan. Gazette, 1896. vol. xxii. p. 293. FITTING, Jahrb. f. wiss. Bot.
10
xxxix. 1903.( ) SACHS, Flora, xlvi. 1863. A. FISCHER, Jahrb. f. wiss. Bot. xxvii.
101 102
1894. PFEFFER, Physiol. Untersuchungen, 1873. ( ) JOST, Bot. Ztg. 1897.
( )
103
MOBIUS, Biol. Zentralbl. 1891. ( ) KLEBS, Biol. Zentralbl. 1889, and Die
Bedingungen der Fortpflanzung bei einigeu Algen und Pilzen, 1896, and Jahrb.
wiss. Bot. 1898 (Sporodinia). OLTMANNS, Morphol. u. Biologic der Algen, vol.
104
ii.
Allgem. Teil, Jena, 1905. ( ) BEYERINCK, Verh. Kon. Akad. v. Wetensch.,
Amsterdam, 1900, p. 352. KLEBS, Jahrb. f. wiss. Bot. xlii. 1906. ( 105 ) v. KOLLIKER,
Ztschr. f. wiss. Zoologie, vol. xiv. 1864, p. 174. H. DE VRIES, Die Mutations-
theorie. Vers. und Beobacht. iiber die Entstehung d. Arten im Pflanzenreich,

Leipzig, Veit and Co. 1901 and 1903. Die Mutationen u. d. Mutationsperioden.
Vortr. a. d. Naturf. - Vers. Hamburg, Leipzig, 1901. Ber. dtsch. Bot. Ges. xxi.
1903. MOLL, Biol. Zentralbl. vol. xxi. 1901, vol. xxii. 1902 (review). KORSCHINSKY,
Flora, 1901, Ergbd. Ixxxix. p. 240. CORRENS, Archiv f. Rassen- und Ges. -Biol.
i. 1904. SOLMS-LAUBACH, Bot. Ztg. Iviii. 1900. JOHANNSEN, see under 119,
DE VRIES, Species and Varieties, their origin by Mutation. Lectures edited by
MAcDouoAL, Chicago and London, 1905. LOTSY, Vorles. iiber Deszendenztheorie,
Jena, 1905.
According to GRAND EURY (Compt. rend, cxlii. 1908) Stigmaria ficoides can
be traced in absolutely constant forms throughout geological strata which must
have required fully a million years for their deposition.
According to ARNOLD ENGLER, characters of the Firs of mountains and plains
are inherited for a long period by their descendants. The same appears to hold
for the Larch and the Alpine Maple. (Mitt. Schweiz. Zentralanst. f. forstl.
Versuchsw. viii. Heft 2, 1905.)

[According to a personal communication Herr Oberfdrster MELSHEIMER found

the form of Shepherd's Purse which SOLMS-LAUBACH named Capsella Hcegeri, in
the year 1882. It occurred in hundreds of individuals on the Dattenberg at Linz
on the Rhine. Specimens still preserved by H. MELSHEIMER are undoubtedly
identical with Capsella Heegeri, Solms.]
106
( ) A. ERNST (Polyembryony), Flora, Ixxxviii. 1901 (includes literature to
1901). FARMER, MOORE and DIGBY, Proc.
LOTSY, Ann. Buitenzorg, 2. Ser. i.
Roy. Soc. vol. Ixxi. 1903. Polyembryonie Euphorbia dulcis HEGELMAIER, v. :

Ber. dtsch. Bot. Ges. xxi. 1903. DE BARY, Bot. Ztg. KUPPER, Flora, xcvi.
KlisTER, Vermehrung und Sexualitat, Vortrag Samml. Aus Natur u. -
1906. :

107
) See
Geisteswelt, Leipzig, 1906. ( literature for Phanerogams (lla). "\\'INKLER
Ber. dtsch. Bot. Ges. xxii. 1905. BITTER, Abh. Nat. Ver. Bremen, xviii. 1904.
108
( LOEB, Amer. Jouru. of Phys. 1900, Vol. iv. p. 178.
) WINKLER, Nachr.
d. Kgl. Ges. d. Wiss., Gottingen, 1900, Heft 2. NATHANSOHN, Ber. deutsch.
Bot. Ges. 1900, p. 99. WINKLER, Jahrb. f. wiss. Bot. vol. xxxvi. 1901, p. 753.
E. STRASBURGER, Bot. Ztg. 1901, ii. Abt. p. 353. JICKELI, Unvollkommenh.
des Stotfwechsels, etc., Berlin, Friedl. Sohn, 1902, and ibid. Vortrag Naturw.
Vcreins, Siebenbiirgen, 1902. KIRCHNER, "Parthenogenesis" in higher plants.
108a
Ber. dtsch. bot. Ges. xxii. 1903. ( ) DIGBY, Proc. Roy. Soc. Ixxvi. 1905.
704 BOTANY

From the j>oiut of view of comparative morphology it is also possible to

speak of apospory in those Hieracia in which an embryo does not develop in the
embryo-sac (macrospore) but (according to O.STK.NKKI.D ami ROSEN ISKKI;) in a cell
of the nucellus which displaces and replaces the embryo-sac.
109 Zeitschr. f. Naturw. xxxvii. 1902, K. F. 30, also as separate
( ) HACKER, Jen.

from G. Fischer. DE VKIES, Befruchtung u. Bastardierung, Leipzig, Veit und

Comp. 1903.
In the Uredineae in which the formation of the secidium starts with a cell
fusion the nuclei remain distinct until the formation of the teleutospores the ;

cells are thus bi-nucleate (Bi-ACKMAX, Ann. of Bot. xviii. 1904. Christman, Bot.
Gaz. xxxix. 1905).
uo Bot. Gaz. 1899, vol. xxviii. pp. 129, 225.
( ) STEVENS, STEVENS, Bot. Gaz.
xxxii. 1901.
A transition to the uninucleate condition of the oospore and to a normal
fertilisation is found in Alb. Tragopogonis.
WINKLER (Merogony), Jahrb. f. wiss. Bot. xxxvi. 1901. (
m ) SOLMS-LAUUA< u,
Bot. Ztg. 1900, ii. 374. STKASBUKGEK, Bot. Ztg. 1900, ii. 307, 1901, ii. 353.
WINKLER, Jahrb. f. wiss. Bot. xxxvi. 1901, 753. ( 112 ) F. KouxiCKE in Handbuch
des Getreidebaues i.
by KORNICKE and WERNER, Berlin, 1885, and Sitzber.
Niederrhein. Ges. f. Natur- u. Heilkunde, Bonn, 1872. NAWASCHIN, Ber.
dtsch. Bot. Ges. 1900, vol. xviii. p. 224. GUIGNARD, Compt. rend. vol. cxxviii.
1899, p. 869, and Ann. sc. nat. Bot., 8. ser. vol. xi. 1900, p. 365, Journ. de Bot.
xv. 1901. DE VRIES, Rev. ge"n. de Bot. vol. xii. 1900, p. 129. CORRENS, Ber.
deutsch. Bot. Ges. 1899, p. 410. STRASBURGER, Bot. Ztg. ii. Abt. 1900, p. 293.
CORRENS. Bastarde zwischen Maisrassen, etc., Bibl. Bot. 1901, Heft 53. ( 11S )
114
CORRENS, Ber. dtsch. Bot. Ges. 1900, vol. xviii. p. 422. ( H. SUHENCK, Die )
11S
Biologie der Wassergewachse, 1886. ( ) CHR. K. SPRENGEL, Das entdeckte
Geheimnis d. Natur im Ban u. in der Befruchtung d. Blume^i, 1793. (Also in
Ostwakl's Klassikern, Xos. 48-51.) DARAVIN, Works. HERM. MULLER, The
Fertilisation of Flowers. F. HILDEBRANDT, Die Geschlechtsverteilung bi-i <lrii

Pflanzen, etc. 1867.

MAC-LEOD, Bot. Jaarb. Dodon. Gent, 1890, gives 688, KNUTH in Handb. d.
Bliitenbiol. to April 1, 1898, 2871 papers on this subject.
FR. JOHOW, Sitzber. K. Pr. Akad. d. Wiss. 1898. VOLKENS, in Schwendener-
Festschrift, 1899. F. JOHOW, Zur Bestiiubungsbiologie Chil. Bliiten, ii. Verh.
d. wiss. Ver. Santiago, vol. iv. p. 345. Valpar. 1901. Includes a list of

Ornithophilous flowers. E. WERTH, Verh. d. bot. Ver. d. Pr., Brandenburg, 1900,

vol. xlii. p. 222. BURCK (Chiropterophilous) flowers, Aim. Buitenzg. 1892. HART,
Bull, of Misc. Inform. 1897. MARLOTH, Ber. deutsch. Bot. Ges. xix. 1901.
DETTO, Flora, xciv. 1905. ANDREAE, Beiheft Bot. Zentralbl. xv. 1903. GILTAY,
Jahrb. f. wiss. Bot. xl. 1904.

Flying insects appear to be mainly attracted by colours, and creeping insects

by scents,but no sharp distinction can be made on these lines.
116
( HILDEBRANDT, Ber. dtsch. Bot. Ges. 1896. Cleistogamy GOBEL, Flora,
)
:

xcv. 1905. Ergbd. HACKEL, Osterr. Bot. Ztschr. Ivi. 1906.

The pollen of chasmogamous flowers also, according to Gbbel, may germinate
within the pollen sacs in a damp hot-house.
Monoecious and Dioecious Mucorineae BLAKESLEE, Proc. of the Amer. Acad.
:

of Arts and Sciences, xl. 4, 1904. Heterostyly ERRERA, Rec. de 1'Inst. bot. :

m
de Bruxelles, 1905. ( ) KOELREUTER, Vorlauf. Nachr. v. einigen das Geschlecht
der Pflanzen betreffenden Versuchen und Beobachtungen, 1761, 1763, 1764, 1766.
 INDEX OF LITERATURE 705

GARTNER, Versuche und Beobachtungen iiber Bastarderzeugung der Pflanzen,

1849. MENDEL, Flora. 1901, Ergbd. Ixxxix. p. 364. CORRENS, Bot. Ztg. 1900,
Sp. 229. HILDEBRANDT, Bot. Zentralbl. vol. Ixxix. 1899, pp. 9, 36. SOLMS-
LAUBACH, Bot. Ztg. 1902, ii. Abt. Sp. 10. TISCHLER, Jahrb. fiir wiss. Bot. xlii.
1906. (
118
) STRASBURGER, Jahrb. fiir wiss. Bot. vol. xvii. 1886. ( ) CORRENS,
m
Sammelref. Bot. Ztg. 1903, ii. 113. Includes reviews of literature. DE VRIF.S,
Compt. rend, cxxxvi. 1903. DE VRIES, Befruchtung u. Bastardierung, Leipzig,
Veit and Comp. 1903. Review Naturw. Rundschau, 1902, p. 640, and Naturw.
Rundschau, 1903, 214. TSCHERMAK, Ztschr. f. d. Landw. Versuchsw. Osterr.
1901, 1902, and 1904. JOHANNSEN, Erblichkeit in Populationen, etc., Jena, G.
Fischer, 1903. CORRENS, Ber. dtsch. Bot. Ges. xx. 1903, and xxi. 1904. CORRENS,
MENDEL'S Briefe an C. NAGELI, Abh. Math. -phys. Klasse Sachs. Ges. d. Wiss.
xxix. 1905. TSCHERMAK, Wien. Landw. Zeitung, 1905. TSCHERMAK, Arch. f.
Rassen- and Gesellsch.-Biologie, 2. LOTSY, Vorles. iiber Deszendenztheorie, Jena,
1905. CORRENS, Vererb. Gesetze, Vortrag, Berlin, 1905. STRASBURGER, Stoffliche
Grundl. der Vererbung, Jena, 1905. STRASBURGER, ALLEN, MIYAKE, and OVERTON,
Histol. Beitrage zur Vererbungsfrage, Jahrb. fiir wiss. Bot. xlii. 1906. MORGAN,
Germ Cells of Mendel pure ? Biol. Zentralbl. Ixxxvi. 1906.
Annual and biennial races also follow Mendel's laws (CoRRENS, Ber. deutsch.
hot. Ges. xxii. 1904).
iao
( ) Zea canina : Contribut. from the Bot. Laborat. Univ. of Pennsylvania, ii.

1901. Cyt. Adami : DARWIN, Ges. Werke, iibers. v. CARUS, 1878, vol. iii.
LAUBERT, Bot. Zentralblatt, Beihefte, x. 1901. TISCHLER, Ber. deutsch. Bot.
Ges. xxi. 1903. Crataegus 1901.
KOHNE, Gartenflora, 1.
:
NOLL, Sitzber.
niederrhein. Ges. f. Natur- und Heilkunde, Bonn, 1905.
STRASBURGER, Jahrb. f.
wiss. Bot. xlii. 1906. (
m
) DINGLER, Ber. dtsch. Bot. Ges. 1887, and Flora, 1887,

and Beweg. der pflanzlichen Flugorgane, 1889. RIDLEY, Ann. of Bot. xix. 1905
(seed). FALCK, Beitr. z. Biol. d. PH., vol. ix. 1904 (spores). ( ) SCHIMPER,
m
Indo-malayische Strandflora, 1891, pp. 158 ff. SCHIMPER, Pflanzengeographie,
123
Jena, G. Fischer, 1900. ( ) BURGERSTEIN, Wien. Gartenztg. 1901. DUGGAR,
Bot. Gaz. xxxi. 1901. SCHULZ, Flora, xi. 1901. Hard seed-coat HILTNER, Arb. :

Biol. Abt. Kaiserl. Ges. -Amt Land- und Forstwirtschaft, 1901, and Naturw.
Zeitschr. fiir Land- und Forstwirtschaft, 1906 (influence of concentrated H 2 S0 4 on
seeds of Conifers). SCHOENE, Flora, xcv. 1905. Erganzungsband.
According to the researches of NEGER (Naturwiss. Zeitschr. fiir Land- und
Forstwirtschaft, vol. ii.) the spores of Bulgaria when in water soon lose their
power of germinating, but this returns on their drying in the vicinity of decaying
parts of plants.
HEINRICHER, Beihefte Bot. Zentralbl. xiii. BORZI, Atti d. Reale
1902. Inga :

124
Acad. d. Lincei Rendiconti, 1903, ser. v. vol. Germination
xii. ( PETER,
) :

Nachr. Getting. Ges. d. Wiss. 1893. SCHINDLER, Die Lehre vom Pflanzenbau auf
physiol. Grundlage, 1896, p. 36 ff. HILTNER, Arb. Biol. Abt. Kais. Ges. -Amt
123
Land- u. Forstwirtschaft, iii. 1902. KLEBS, Unters. Bot. Inst. Tubingen, vol.
i.
p. 536 S. (
m
) PFITZER, Ber. deutsch. Bot. Ges. 1885. (
(
127
)

) Cucurbita cf. 88. :

DE VRIES, Landw. Jahrbiicher, 1880. RIMBACH, Die kontraktilen Wurzeln.

FUNFSTUCKS, Beitr. z. wiss. Bot. vol. ii. 1897, and Ber. dtsch. Bot. Ges. 1899,
vol. xvii. p. 18.

2 z
706 BOTANY

CRYPTOGAMS

KLEBS, Die Beding. der Fortpflanz. bei niederen Algen und Pilzen, 1896, and
l
( )

Jahrb. f. wiss. Botanik, xxxiu-xxxiv. further Willkiirl. Entwicklgsand. bei Pflanzen,

;

1903. ( 2 ) A. FISCHER, Vorlesungen iiber Bakterien, 1897, 2nd edition, 1903. FLUGGE,
Die Mikroorganismen, 1896. MIGULA, System der Bakterien, 1897-1900, and Nat.
Pflfam. I 1 MIGULA, A. de Bary's Vorlesungen iiber Bakterien, 3rd edition, 1900.
.

KOLLE and WASSERMANN, Handb. der pathogen. Mikroorganismen, 1904. A. M KVKII,

3
Flora, 1899, p. 428. ( ) SORAUER, LINDAU, REH, Handbuch der Pflanzenkrank-
4
heiten, 1906. ( ) THAXTER, Bot.
Gaz. xiv. 1892, p. 389, xxiii. 1897, p. 395, and
xxxvii. 1904, p. 405. QUEHL, Ctrbl. fur Bakt., Parasitenkunde, etc., xvi. 1896,
5
p. 9. KIRCHNER, in Nat. Pflfam. I
( )
1
A. FISCHER, Unters. iiber den Bau der .

Cyanophyceen und Bakterien, 1897 also Bot. Zeitg. 1905, p. 51. ZACHARIAS,
;

Abhand. aus dem Geb. der Naturw., Hamburg, 1900, and Jahrb. d. Hamburger
wiss. Anstalten, xxi. 1903. HEGLER, Jahrb. f. wiss. Bot. xxxvi. 1901, p. 229.
MASSART, Recueil de 1'inst. bot. de Bruxelles, v. 1902. BRAND, Ber. deutsch.
bot. Ges. 1901, p. 152, 1905, p. 62, and Beihefte bot. Zentralbl. xv. 1903, p. 31.
KOHL, Organisation und Physiol. d. Cyanophyceenzelle, Jena, 1903, and Beihefte
bot. Zentralbl. 1905, xviii. p. 1. FRITSCH, Beihefte bot. Zentralbl. 1905, xviii.
p. 194. OLIVE, Beihefte bot. Zentralbl. 1905, xviii. p. 9. PHILIPPS. Contrib. bot. Lab.
6
Pennsylvania, ii. 1904, p. 237. ( ) OLTMANNS, Morphologic und Biologie der Algen, i.
7 8
1904. ( ) SENN, in Nat. Pflfam. I 1 ( ) ZUMSTEIX,
. Jahrb. f. wiss. Bot. xxxiv.
1900, p. 149.
9
( ) SCHROTER,
in Nat. Pflfam. I 1 DE BARY, Comparative Morph. .

and Biol. of the Fungi, Mycetozoa and Bacteria. ZOPF, Die Pilztiere, 1885.
LISTER, A Monograph of the Mycetozoa, 1894. HARPER, Bot. Gaz. xxx. 1900, p.
10
217. JAHN, Ber. deutsch. bot. Ges. 1904, p. 84, 1905, p. 489. ( ) WORONIN,
Jahrb. wiss. Bot. xi. 1878, p. 548.
f. NAWASCHIN, Flora, 1899, p. 404. PROWAZEK,
Arb. kais. Gesundheitsamt, xxii. 1905, p. 396. n SCHUTT, in Nat. Pflfam. I 1 ,
( )

and Die Peridin. der Planktonexpedition, 1895. SCHILLING, Flora, 1891, p. 220.
12
( ) SCHUTT, Das Pflanzenleben der Hochsee, 1893. GRAN, Das Plankton des
13
norwegischen Nordmeeres, 1902. ( ) SCHILLING, Ber. deutsch. bot. Ges. 1891,
199. M ZEDERBAUER,
Ber. deutsch. bot. Ges. 1904, p. 1. 15
DE BARY,
p. ( ) ( )

Unters. iiber die Konjugaten, 1858. RALFS, The British Desmidieae, 1848. WILLE,
in Nat. Pflfam. P. W. WEST and G. S. West, A Monograph of the Brit. Desmid.
i. 1004. LiiTKEMULLER, Beitr. z. Biol. d. Pflanz. viii. 1902, p. 347. KLEBAHN,
16
Jahrb. f. wiss. Bot. xxii. 1891, p. 415. ( ) CHMIELEWSKY, Arb. d. Gesellsch. d.
17
Naturf. d. Charkower Universitat, xxv. 1890. ( ) SMITH, Synopsis of the Brit.
Diatom. 1853-56. SCHMIDT, Atlas der Diatomeenkunde, Leipzig, 1874-77. VAN
HEURCK, Synopsis des Diat. 1880-85, and Traite des Diat., Anvers, 1899. DIPPEL,
Diatomeen der Rhein-Mainebene, 1905. PFITZER, in Bot. Abh. von Hanstein I 2 ,

1871. KLEBAHN, Jahrb. f. wiss. Bot. xxix. 1896, p. 595. KARSTEN, Flora, 1896,
p. 286, 1897, pp. 33, 203, 1900, p. 253, and Die Diatomeen der Kieler Bucht, 1899.
SCHU'TT, in Nat. Pflfam. i., and Jahrb. f. wiss. Bot. 1899-1900. 0. MULLER, Ber.
18
deutsch. bot. Ges. 1898-1903. ( ) GRAN,
Die Diat. der arkt. Meere, Fauna arctica,
iii. KARSTEN, Ber. deutsch. bot. Ges. 1904, p. 544, and Wiss. Ergebn. der
1904.
19
d. Tiefseeexped. ii. 1905. ( ) BENECKE, Jahrb. f. wiss. Bot. xxxv. 1900, p. 535.
20
KARSTEN, Flora, Ergzb. 1901, p. 404. ( ) KCTZING, Tabulae phycologicae.
(
21
) WILLE, in Nat. Pflfam. I
2
(
M ) WILLE, Algolog. Notizen ix.-xiv., Christiania,
.

1903. m
SCHMIDLE, Ber. deutsch. bot. Ges. 1903, p. 346. ( ) GOROSCHANKIN,
 INDEX OF LITERATURE 707

Flora, 1905, p. 420. (

M ) KLEIN, Jahrb. f. wiss. Bot. xx. 1889, p. 133, and Ber. nat.
Ges. Freiburg, v. 1890. OVERTON, Bot. Zentralbl. xxxix. 1889, p. 65. A. MEYER,
25 26
Bot. Ztg. 1896, p. 187. O ) BEYERINCK, Bot. Ztg. 1890, p. 720. f ) GRINTZESCO,
27
Rev. gener. de hot. xv. 1903, p. 5. ( ) SENN, Bot. Ztg. 1899, p. 39. ()
29
ASKENASY, Ber. deutsch. hot. Ges. 1888, p. 129. ( ) KLEBS, Bot. Ztg. 1891,
p. 789, and Flora, 1890, p. 351. TIMBERLAKE, Transact. Wiscons. Acad. xiii.
30
and Annals of Bot. xv. 1901, p. 619. DODEL, Jahrb. f. wiss. Bot. x. 1876, ( )

31
p. 417. KLEBS, Beding. der Fortpfl. bei Algen und Pilzen, 1896. PRINGSHEIM, ( )

Jahrb. f. wiss. Bot. i. 1858, p. 1. JURANYI, ibid. ix. 1873-74, p. 1. KLEBAHN,

32
ibid. xxiv. 1892, p. 235. ( ) PRINGSHEIM, ibid. ii. 1860, p. 1. OLTMANNS,
ALLEN, Ber. d. bot. Gesellsch. 1905, p. 285. C ) DE BARY
33
Flora, 1898, p. 1.

and STRASBURGER, Bot. Ztg. 1877, p. 713. SOLMS-LAUBACH, Trans. Linn.

Soc. 1895. (*) REINKE, Wiss. Meeresunters. Kiel, 1899. WEBER VAN BOSSE, ,

Annales jard. Buitenzorg, xv. 1898, p. 243. HABERLANDT, Sitzb. Akad.

Wien, cxv. i. 1906, p. 1. p) ERNST, Beiheft bot. Zentralbl. xiii. 1903, p. 115.
OLTMANNS, Flora, 1895, 388. x
DAVIS, Bot. Gazette, xxxviii. 1904, p. 81. ( )
p.
2
WILLE, in Nat. Pflfam. I .
MIGULA, in Rabeuhorst, Krypt. Flora Deutschland,
1890. GIESENHAGEN, Flora, 1896, p. 381, and 1898,
GOTZ, Bot. Ztg. 1899, p. 1.

p. 19. GOEBEL, MOTTIER, Annals of Bot. xviii. 1904, p. 245.

Flora, 1902, p. 279.
37 2
( ) KJELLMANN, in Nat. Pflfam. I POSTELS et RUPRECHT, Illustr. algarum .

Oceani pacifici, 1840. HOOKER, Flora antarctica, i. 1844. REINKE, Atlas deutscher
Meeresalgen, 1889-92. REINKE, Studien zur vgl. Entwicklgsgesch. der Lami-
narien, Kiel, 1903.
M ) SKOTTSBERG, Bot. Notiser, 1903, 39
HANSTEEN,
( p. 40. ) (

Jahrb. f. wiss. Bot. xxxv. 1900, p. 611. HUNGER, ibid, xxxviii. 1903, p. 70. (
w )
41
WILLE, Univers. Festschrift, Christiania, 1897. ( ) BERTHOLD, Flora, 1897, p. 398.

OLTMANNS, Flora, 1897, p. 398 ; 1899, p. 86. SAUVAGEAU, Journ. de botan. 1896.
42
( ) CHURCH, Annals of Bot. xii. 1898, p. 75. SAUVAGEAU, Ann. sc. nat. 8 ser. x.
43
1899, p. 265. t ) WILLIAMS, Annals of Bot. xi. 1897, p. 545, and xviii. 1904,
pp. 141, 183 xix. 1905, p. 531.
; (**) MOTTIER, Annals of Bot. xiv. 1900, p. 163.

STKASBURGER, Bot. Ztg. 1906, ii. p. 3. ( ) THURET and BORNET, Etud. phycolog.
45

1877. OLTMANNS, Bibl. bot., Heft 14. STRASBURGER, Jahrb. f. wiss. Bot. xxx.
1897, p. 351. FARMER and WILLIAMS, Phil. Trans. Roy. Soc. London, vol. cxc.
46 2
1898. ( ) SCHMITZ and HAUPTFLEISCH, Nat. Pflfain. I FALKENBERG, xxvi. .

Monogr. Fauna und Flora, Neapel, 1901. KOLKWITZ, Wiss. Meeresunters., Kiel,
1900. WILLE, Ber. deutsch. bot. Ges., 1894, p. 57, and Nova Acta, Hi., 1887, p. 51.
OSTERHOUT, Flora, 1900, p. 109. OLTMANNS, Bot. Ztg. 1898, p. 99. HASSENKAMP,
Bot. Ztg. 1902, p. 65. WOLFE, Annals of Bot. xviii. 1904, p. 607. YAMANOUCHI,
47
Bot. Gazette, xli. 1906, p. 425. ( ) KUCKUCK, Sitzb. Akad. Berlin, 1894, p. 983.
STURCH, Annals of Bot. xiii. 1899, p. 83. C ) DE BARY, Comp. Morph. and
48

49
Biology of the Fungi. ( ) BHEFELD, Bot. Unters. iiber Schimmelpilze, Unters.
aus dem Gesamtgebiet d. Mykologie, 1872-1905. VON TAVEL, Vgl. Morphol. d.
50 51 1
Pilze, 1892. ( ) SCHROTER,
in Nat. Pflfam. I ( ) CORNU, Ann. sc. nat., 1872. .

THAXTER, Bot. Gaz. 1895. LAGERHEIM, Bihang till Svensk. Akad. Handl. xxv.
WORONIN, Mem. de 1'Acad. imp. des Sciences de St-Petersbourg, 1904, 8. ser.
52
vol. xvi. (
No.) TROW, Annals of Bot. ix. 1895, p. 609 ; xiii. 1899, p.
4, p. 1.
130 xviii. 1904, p. 541. KLEBS, Jahrb. f. wiss. Bot. xxxiii. 1899, p. 513. DAVIS,
:

53
Bot. Gaz. xxxv. 1903, p. 233. ( ) WORONIN, Jahrb. f. wiss. Bot. xi. 1978, p. 556.
M ) WAGER, Annals of Bot. iv. 1889-91, 127 x. 1896, pp. 89, 295 xiv.
( p. ; ; 1900,
p. BERLESE, Jahrb. f. wiss. Bot. xxxi. 1898, p. 159. DAVIS, Bot. Gaz.
263.
xxix. 1900, p. 297. STEVENS, Bot. Gaz. 1899, xxviii. p. 149 1901, xxxii. p. 77 ; ;

1902, xxxiv. p. 420, and Ber. deutsch. bot. Ges. 1901, p. 171. TROW, Annals of
708 BOTANY

Bot. xv. 1901, p. 269. MIVAKE, Annals of Bot. xv. 1901, p. 653. ROSENBERG,
Bihang till Svensk. Ak. vol.' xxviii. 1903. Krai, AND, Jahrb. f. wiss. Bot. xxxix.
1904, p. 135. ROSTOWZEW, Flora, 1903, p. 405. (
M ) HARPER, Annals of Bot. xiii.

1899, p. 467. GIIUBER, Ber. deutsch. bot. Ges. 1901, p. 51. DANGEARD, Conipt.
rend. acad. 1906, p. 645.
Paris, cxlii. (
x ) BLAKESLEE, Proceed. Amer. Acad.
xl. 1904. (
67
)
M
OLIVE, Bot. Gaz. xli. 1906, pp. 192, 229. ( ) EIDAM, Cohn's Beitr.
z. Biol. iv. 1887, p. 181. RACIBOUSKI, Flora, 1906, p. 106. FAIRCHILD, Jahrb. f.
wiss. Bot. xxx. 1897, p. 285. WovcrcKi, Flora, 1904, p. 87. (
w TULASXE,
) Selecta
Fungorum Carpologia, 1861-65. SCHROTER, LIXDAU, FISCHER in Nat. Pflfam. I 1
.

w ) HARPER, Ber. deutsch. bot. Ges. 1895,

( p. 67, and Jahrb. f. wiss. Bot. xxx.
1897, p. 249 also Annals of Bot. xiii. 1899, p. 467
;
xiv. 1900, p. 321. GUILLIER- ;

MOND, Revue gener. de bot. xvi. 1904, pp. 49, 130. CLAUSSEX, Bot. Ztg. 1905,
) DE BARY, Beitr. z. Morphol. und Phys. der Pilze, iii. 1870.
61
p. 1. ( HARPER,
Ber. deutsch. bot. Ges. 1895, p. 475, and Jahrb. f. wiss. Bot. xxix. 1895, p. 655.
NEGER, Flora, 1901, p. 333, and 1902, p. 221. SALMON, Annals of Bot. xx. 1906,
p. 187. HARPER, Carnegie Institution of Washington, publ. No. 37, 1902
(Phyllactiniaj. (**) HARPER, Annals of Bot. xiv. 1900, p. 321 (Pyroueuia).
CLAUSSEN, Bot. Ztg. 1905, p. 1 (Boudiera). (*) KROMBHOLZ, Abb. und Beschreib.
der Schwamme, 1831-46. LENZ, Niitzl., schiidl. und verdacht. Schwiininie, 1890.
SCHLITZBERGER, Pilzbuch. MICHAEL, Fiihrcr fur Pilzfreunde. ROLL, Unsere
essb. Pilze. ( M ) GOETHE, Uber den Krebs der Obstbiiume, Berlin, 1904. ( M ) FISCUER
66
in Rabenhorst, Krypt. Flora, I 5 ) SADEBECK, Jahrb. Hamburg, wiss. Anst. 1884,
.
(

1890, 1893, and Ber. d. bot. Ges. 1903, p. 539 1904, p. 119. GIESENHAGEX,;

Flora, Ergzb. 1895, p. 267, and Bot. Ztg. 1901, p. 115. IKENO, Flora, 1903, p. 1.
(
w ) GUILLIERMOND, Rev. gener. de bot. 1903, p. 337. THAXTER, Mem. of Amer.
) DIETEL, LINDAU, HEXX-
1 70
Acad., Boston, 1896. LIXDAU, in Nat. Pflfam. I ( .

1 71
INGS, FISCHER in Nat. Pflfam. I ( ) JUEL, Jahrb. f. wiss. Bot. xxxii. 1898,
.

p. 361. RUHLAXD, Bot. Ztg. 1901, p. 187. HARPER, Bot. Gaz. xxxiii. 1902, p. 1.

C
72
HARTIG, Krankh. der Waldbaume, 1874, and Lehrb. der Pflauzenkrankheiten,
) R.
1900. (
73
) HARPER, Transact. Wisconsin Acad. 1899, p. 475. ( 74 ) BREFELD and
FALK, tint, aus dem Gesammtgeb. d. Mykologie, xiii. 1905. HECKE, Ber. d. Bot.
) Numerous papers, MAGNUS, KLEBAHX, SYDOW,
Ges. 1905, p. 248. (
7:>
ERIK.-MIN.
76
TISCHLER, E. FISCHER, LAGERHEIM, and others. ( ) BLACKMAX, Annals of Bot.
xviii. 1904, p. 323. BLACKMANX and FRASER, ibid. xx. 1906, p. 35. CHRISTMAN.
77
Bot. Gaz. xxxix. 1905, p. 267. ( ) KLEBAHX, Die wirtswechselnden Rostpilze,
1904. (
ra
) ERIKSSON and TISCHLER, Svenska Vet. Akad. Handl. 1904, vol. xxxvii.,
xxxviii. Ber. deutsch. bot. Ges. 1904, p. 255.
KLEBAHN, E. FISCHER, Bot. Ztg.
1904, p. 327. MARSHALL WARD, Annals
of Bot. xix. 1905, p. 1. 79
C ) MOLISCH,
Leuchtende Pflanzen, 1904. O
R. HARTIG, Der echte Hausschwamm, 1885 ; 2nd
vox TUBETF, 1902. (") MOLLER, Pilzgarten feiidamerik. Ameisen, 1893.
edition by
HOLTERMANX, Schwendener Festschrift, 1899. FOREL, Biolog. Zentralbl. 1905, p.
82
170. HUBER, Biolog. Zentralbl. 1905, p. 606. C E. FISCHER, Denkschr. Schweiz. )
M
nat. Ges. xxxii. and xxxvi. MOLLER, Brasil. Pilzblumen, 1895. ( ) FUXFSTUCK
and ZAHI.BILUCKXER, in Nat. Pflfam. I REIXKE, Jahrb. f. wiss. Bot. 1894-96.
1
.

SCHWEXDEXER, Algentypen der Flechtengonidien, 1869. BOXXIER, Ann. sc. nat.

1889. BITTER, Hedwigia, xl. 1901, p. 171, and Jahrb. f. wiss. Bot. xxxvi. 1901,
M ) PEIRCE, Proceed. Calif. Acad. 1899, p. 207, and The Amer. Naturalist,
p. 421. (
85
1900, p. 245. ELENKIN, Bull. Jard. Bot. St-Petersb. iv. 1904. C ) ZOPF, Beih.
86 87
bot. Zentralbl. xiv. 1903, p. 95. C ) STAHL, Hiickelfestschrift, 1904, p.
357. C )

1891. Bot. 21. STAHL, Beitr. z. Entw.

KRABBE, Cladonia, BAUR, Ztg. 1904, p. (<*)

d. Flechten, 1877. BAUR, Ber. deutsch. bot. Ges. 1898, p. 363, and Flora, 1901,
 INDEX OF LITERATURE . 709

p. 319 ;
Bot. Ztg. 1904, p. 21. DARBISHIRE, Jahrb. f. wiss. Bot. xxxiv. 1900,
p. 329. LINDAU, Flora, 1888, p. 451, and Schwendenerfestschrift, 1899. WOLFF,
89
Flora, Ergzsb. 1905, p. 31. ( ) HOLLER, Kultur flechtenbild. Ascomyceten, 1887 ;

Bot. Ztg. 1888, p. 421. (

w ) GLUCK, Flechtenspermogonien, Habilitationsschr.
91
Heidelberg, 1899. ) JOHOW, Jahrb. f. wiss. Bot. xv. 1884, p. 361.
( HOLLER,
92
Flora, 1893, p. 254. POULSEN, Vid. Hedd. Kopenhagen, 1899. t ) GOEBEL, Die
Huscineen. Schenks Hdb. d. Bot. ii. 1882 Organography, vol. ii. Flora, 1902. ; ;

(
m ) CAMPBELL, The Structure and Development of Hosses and Ferns, 2nd edition,
94
1905. ( ) GOEBEL, Flora, 1902, p. 279. DAVIS, Annals of Botany, xvii. 1903, p.
477. HOLFEKTY, Bot. Gaz. xxxvii. 1904, p. 106. ( 9S ) IKENO, Beih. bot. Zentralbl.
xv. 1903, p. 65 (Harchantia). JOHNSON, Bot. Gaz. xxxviii. 1904, p. 185 (Hono-
clea). LEWIS, Bot. Gaz. xli. 1906, p. 110 (Riccia). BOLLETER, Beih. bot. Zentralbl.
xviii.
1
1905, p. 327 (Fegatella). (
M ) PFEFFER, Unters. bot. J. Tiibingen, i., ii.
Bot. 97 in Nat. Pflfam.
LIDFORS, Jahrb. f. wiss. xli. 1904, p. 65. ( ) SCHIFFNER,

P. LEITGEB, Unters. iiber GOEBEL, Flora, 1895, p. 1.

Lebermoose, 1874-82.
98
ANDREAS, Flora, 1899, p. 161. GARBER, Bot. Gaz. xxxvii. 1904, p. 161. LEWIS,
C )

Bot. Gaz. xli. 1906, p. 110. (") KNY, Botan. Wandtafeln, viii. 1890. IKENO,
Beih. bot. Zentralbl. xv. 1903. KAMERLING, Flora, Ergzb. 1897, p. 1. BOLLETER,
Beih. bot. Zentralbl. xviii. 1 1905, p. 327. ( 10 ) GOTTSCHE, Haplomitrium, Nova Acta
Acad. Leop. xx. 1843, p. 267. GOEBEL, Ann. jard. Buitenzorg, ix. 1890, p. 11,
101
and Organography, vol. ii. p. 39. ( ) K. HULLER, RUHLAND, WARNSTORF,
BROTHERUS in Nat. Pflfam. P. LORENTZ, Jahrb. f. wiss. Bot. vi. 1867-68, p. 363.
HABERLANDT, ibid. xvii. 1886, p. 359. TANSLEY and CHICK, Annals of Bot. xv.
1901, p. 1. CORRENS, Vermehrung der Laubmoose, 1899. VAUPEL, Flora, 1903
102 103
p. 346. ( ) ZEDERBAUER, Ostr. bot. Ztschr. 1 902. ( ) HABERLANDT, Jahrb. f.

wiss. Bot. xvii. PORSCH, Der Spaltbffnungsapparat im Lichte der

1880, p. 357.
Phylogenie, 1905, p. 33.
104
PH. W. SCHIMPER, Torfmoose, 1858. NAWASCHIN,
( )
105
Flora, 1897, p. 151. ( ) LANTZIUS-BENINGA, Der innere Bau der Mooskapsel, 1850.

DIHM, Flora, Ergzbd. 1894, p. 286. GOEBEL, Flora, 1895, p. 459. STEINBRINCK,
106 4
Flora, Ergzbd. 1897, p. 131. ) SADEBECK, in Nat. Pflfam. I GOEBEL, Organ-
( .

2 107
ography II ( ) PFEFFER, Unters.
. bot. Inst. Tubingen, i. p. 363 (Ferns, Sela-
ginella). SHIBATA, Bot. Hag. Tokyo, xix. 1905, p. 39 (Salvinia) ibid. pp. 79, 126 ;

(Equisetum) Ber. d. bot. Ges. 1904, p. 478, and Jahrb. f. wiss. Bot. xli. 1905,
;

10B
p. 561 (Isoetes). LIDFORS, Ber. d. bot. Ges. 1905, p. 314 (Equisetum). ( )

SADEBECK, DIELS, BITTER in Nat. Pflfam. I 4 HOOKER, Synopsis filicum, 1883. .

BAKER, Fern Allies, 1887. CHRIST, Farnkrauter der Erde, 1897. BOWER,
Trans. Roy. Soc. 1899, and Proc. Roy. Soc. 1903. 109
( ) JEFFREY, Univers. of
Toronto, Biolog. Series, No. 1, 1898 (Botrychium). BRUCHMANN, Bot. Ztg. 1904,
p. 1 (Ophioglossum), and Flora, 1906, p. 203 (Botrychium). CARDIFF, Bot. Gaz.
xxxix. 1905, p. 340. (
uo
) LYON, Bot. Gaz. xl. 1905, p. 455 (Botrychium). ( )
m
STEINBRINCK, Ber. deutsch. bot. Ges. 1903, p. 217. URSPRUNG, Jahrb. f. wiss. Bot.
xxxviii. 1903, p. 635, and Ber. deutsch. bot. Ges. 1904, 112
p. 73. ( ) STRASBURGER,

Jahrb. f. wiss. Bot. vii. 1869-70, p. 390. (

113
) DE BARY, Bot. Ztg. 1878, p. 449.
BOWER, Trans. Linn. Soc. 1887. LANG, Bot. Zentralbl. Ixxiii. 1898, p. 145.
FARMER, HOORE, DIGBY, Proc. Roy. Soc. Ixxi. 1903. (
m) PRINGSHEIM,
Jahrb.
wiss. Bot. 116
f. 1863, p. 484. BELAJEFF, Bot. Ztg. 1898, p. 141. ( ) STRAS-
iii.

BURGER, Azolla, 1873. ( 116 ) SADEBECK, in Nat. Pflfara. I 4 JEFFREY, Hem. of .

Boston, Soc. of Nat. Hist. v. URSPRUNG, Jahrb. f. wiss. Bot. xxxviii. p. 655 ; Ber.
deutsch. bot. Ges. 1904, p. 73. STEINBRINCK, Ber. deutsch. bot. Ges. 1903, p. 217.
(
m ) URSPRUNG, Jahrb. f. wiss. Bot. xxxviii. p. 635. (
118
) PRITZEL, in Nat. Pflfam.
I 4. GOEBEL, Bot. Ztg. 1887, p. 161. TREUB, Ann. Jard. Buitenzorg, 1884-90.
 710 BOTANY

BRUCHMAXN, Uber die Prothallien und Keimpfl. der Lycop. 1898. LAXG, Annals
119 4
of Bot. 1899, p. 279.
xiii. ( ) HIERONYMTS, in Nat. Pilfam. I PFKFFF.K, in
.

Hansteins Bot. Abh. 1871. BELAJEFF, Bot. Ztg. 1885, p. 793. BRITHMANN,
Unters. iiber Selaginella spinulosa, 1897. FITTING, Bot. Ztg. 1900, p. 107. GOEBEL,
Flora, 1901, p. 207. LYON, Bot. Gaz. xxxii. 1901, p. 124, and xl. 1905, p. 285.
CAMPBELL, Annals of Bot. xvi. 1902. p. 419. DEXKE, Beiheft z. Bot. Zentralbl. xii.
12
1902, p. 182. STEIXBRIXCK, Ber. deutsch. bot. Ges. 1902, p. 117. ( ) BRUCH-
121
MAXX, Flora, 1905, p. 150. GOEBEL, Flora, 1905, p. 195. ( ) HABERLANDT, Flora,
122
1888, p. 291, and Ber. d. bot. Ges. 1905, p. 441. ( ) SADEBECK,
in Nat. Pflfam.
I4.
FARMER, Annals of Bot. v. 1890-91, p. 37. SCOTT and HILL, ibid. xiv.
1900, p. 413. SMITH, Bot. Gaz. xxix. 1900, p. 225. FITTING, Bot. Ztg. 1900,
p. 107. SOLMS - LAUBACH, ibid. 1902, p. 178. BELAJEFF, ibid. 1885, p. 793.
CAMPBELL, Mosses and Ferns, p. 536. ( ) W. PH. SCHIMPER, Traite de Paleon-
123

tologie vege"tale, Paris, 1866-1874. W.

PH. SCHIMPEU and A. SCHEXK, Palaeo-
phytologie, in Zittel's Handbuch Pal. 1890. A. SCHEXK, Die fossilen
der
Pflanzenreste, 1888. B. RENAULT, Cours de bot. fossile, 1881-85. G. SAPOUTA
and MARION, Evolution du regne vegetal, 1881 and 1885. SOLMS-LAUBACH, Fossil
Botany. POTONIE, Lehrb. der Pflanzenpaliiontologie, 1899. D. H. SCOTT, Studies
4
in Fossil Botany, 1900. POTONIE, Fossile Pteridophyten in Nat. Pflanzenfam. I .

R. ZEILLER, Elements de paleobotanique, 1900.

PHANEROGAMS
l
( )
W. HOFMEISTER, Vergleich. Untersuch. der Keim., Entfalt. und Fruchtbild.
hoherer Kryptogamen und der Samenbild. der Koniferen, Leipzig, 1851. ( 2 ) E.
STRASBURGER, Angiospermen und Gymnospermen, Jena, 1879. OVEUTOX, Reduk-
tion der Chromosomen, Vierteljahrsschr. d. natnrf. Ges. Zurich, 1893. JUEL,
Tetradenteilg., Jahrb. f. wiss. Bot. xxxv. 1900. M. KOERNICKE, Niederrhein. Ges.
1901. COULTER and CHAMBERLAIN, Morphology of Spermatophytes I., New York,
1901. K. GOEBEL, Organography of Plants. E. WARMING, Pollenbildende
Phyllome und Caulome, Hanst. bot. Abh. ii. 1873. STRASBURGER, Bau und
Wachstum d. Zellhaute, 1882. Wachstum vegetab. ZellhJiute, 1889. Befruchtungs-
vorgang d. Phanerogamen, Jena, 1884. WILLE, Pollenkorner d. Angiospermen,
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Tubingen, i. 1884. (**) E. STRASBURGER, Reduktionsteiltmg. Sitzgsber. K. Akad.
d. W. Berlin, xviii. 1904. STRASBURGER, CH. ALLEX, KIICHI MIYAKE, I. BERTH.
OVERTOX, Histolog. Beitr. Jahrb. f. wiss. Bot. xlii. Leipzig, 1905. STRASBURGER,
,

Stoffl. Grundlagen der Vererbung, Jena, 1905. 0. ROSENBERG, Reduktionsteilung.

Botau. Notiser, 1905. ( 3 ) EICHLER, Bliitendiagramme i. and ii. Leipzig, 1875, 1878.
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ii. iii. iv. from 1889. BAILLON, Histoire des plantes i.-xiii. 1867-94. BERG
and SCHMIDT, Atlas d. offizin. Pflanzen, 1863, and the second edition by A. MEYER
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W. HOFMEISTER, Vgl. Untersuchungen, 1851.
( )

STRASBURGER, Befruchtg. bei d. Koniferen, Jena, 1869. Koniferen und Gnetaceen,

Jena, 1872. Angiospermen und Gymnospermen, Jena, 1879. COULTER and
CHAMBERLAIN, Morph. of Spermatophytes, i. and ii., N.Y. 1901 and 1903.
GOEBEL, Organography of Plants. The literature cited in 3 and 4 includes the
 INDEX OF LITERATURE 711

most important and fundamental works dealing with the Phanerogams as a whole ;

5
these works will not be referred to below. ( ) L. CL. RICHARD, Comrnentatio
de Coniferis et Cycadeis, Stuttgart, 1826. E. WARMING, Cycadeerne, K. D.
Vidensk. Selsk. Forh. 1877 and 1879. M. TREUB, Cycadees, Ann. de Buitenzorg,
2 and 4, 1885. W. H. LANG, Stangeria, Ann. of Botany, xi. 1897, and xiv. 1900.
H. J. WEBBER, Spermatogenesis and Fecundation of Zamia, Washington, 1901,
U.S. Dep. of Agricult. S. IKENO, Cycas revoluta, Jahrb. f. wiss. Bot. xxxii. 1898.
H. Graf SOLMS- LAV BACH, Stangeria, Bot. Ztg. 1890. G. KRAUS, Ann. de
Buitenzorg, xiii. 1896. COULTER and CHAMBERLAIN, Zamia, Bot. Gaz., March
1903. M. C. STOPES, Cycadeen, Flora, 1904, 435. H. MATTE, Cycadacees, Caen,
1904. H. MIYAKE, Cycas -Spermatozoiden, Ber. der deutschen botan. Gesellsch.
1906. 6
( )
Literature 4, and SAKUGORO HIRASE, Ginkgo biloba, Botan. Zentralbl.
vol. Ixix. No. 2, 1897. HIRASE, Journ. of the College of Science, Univ. Imp. Tokio,
viii. 1895, and xii. 1898. K. GOEBEL, Pollenentleerung, Flora, 1902, Ergbd.
7
( ) and GOROSCHANKIN, Corpuscula, Bot. Ztg. 1883. J. SACHS,
Literature 4,
Lehrb. d. Botanik, iv. 1874. L. CELAKOWSKY, Gymnospermen, Abh. kg. bohm.
Ges. d. W. 1890. Continued in Engler's Jahrb. 1898. BELAJEFF, Ber. d.
deutsch. bot. Ges. 1891, 280, and 1893, 196. DIXON, Pinus silvestris, Ann. of Bot.
1894. NOLL, Abietineen-Zapfen, Sitzgsber. d. niederrh. Ges. 1894. JAGER, Taxus
baccata, Flora, 1899. ARNOLDI, Gymnospermen, 1, 2, and 5 Bull, de Moscou, 1899,
1900 3 and 4 Flora, 1900, 6 Russian with resume in German
;
J. WIESNER, ;

Rohstoffe des Pflanzenreichs, 2nd edition, 1900. GOEBEL, Pollenentleerung, Flora,

Ergbd. 1902. K. FUJII, Bestaubungstropfen, Ber. d. deutsch. Bot. Ges. 1903.
F. W. OLIVER, Older Gymnosperms, Ann. of Bot. xvii. 1903. K. MIYAKE, Abies
balsamea, Beih. z. bot. Zentralbl. 1903. MIYAKE, Picea excelsa, Ann. of Bot.
xvii. 1903. A. LAWSON, Sequoia, Ann. of Bot. xviii. 1904. LAWSON, Cryptomeria,
Ann. of Bot. xviii. 1904. E. STRASBURGER, Taxus, Haeckel-Festschrift, Jena,
1904. JUEL, Cupressus, Flora, 1904. W. C. COKER, Taxodium, Bot. Gaz. July and
Aug. 1903. COKER, Bot. Gaz. 38, 1904. MARGARET FERGUSON, Pinus, Proc.
Washington Acad. of Sc., vi. Oct. 1904. COULTER and LAND, Torreya, Bot. Gaz.
xxxix. 1905. E. STAHL, Mexik. Nadelhblzer and L. KLEIN, Mitteleurop. Waldbaume
in " Vegetationsbilder," ii. Reihe, 3 and 5-7, Jena, 1904. 8
( ) Literature 4 and
J.D. HOOKER, Welwitschia, etc., Trans. Linn. Soc., Lond. xxiv. 1863. F. 0.
BOWKR, Welwitschia, Quart. Journ. Micr. Soc. xxi. 1881. BOWER, Gnetum Gnemon,
ibid. xxii. 1882. H. H. W. PEARSON, Welwitschia, Phil. Trans. Roy. Soc.

Lond., vol. cxcviii. KARSTEN, Gnetum, Bot. Ztg. 1892.

1906. G. KARSTKX,
Ann. de Buitenzorg, xi. 1893. KARSTEN, Cohn's Beitr. z. Biol. vi. 1893. JACCARD,
Ephedra helvetica, Diss., Zurich, 1893. LOTSY, Gnetum, Ann. de Buitenzorg, xvi.
1899. LOTSY, Parthenogenesis, Flora, 1903. W. J. G. LUND, Bot. Gaz. xxxviii. 1904.
9
( W. PH. SCHIMPER, Traite de Pale"ontologie vegetale, Paris, 1866 74.
)
-

SCHIMPER and A. SCHENK, Palaeophytologie in Zittel's Handbuch der Pal. 1890.

A. SCHENK, Die fossilen Pflanzenreste, 1888. B. RENAULT, Cours de bot. fossile
1881-85. G. SAPORTA and MARION, Evolution du regne vegetal, 1881 and 1885.
H. GRAF zu SOLMS-LAUBACH, Fossil Botany. POTONIE, Lehrbuch der Pflanzen-
palaontologie, 1899 D. H. SCOTT, Studies in Fossil Botany, 1900 R. ZEILLER,
; ;

Elements de paleobotanique, 1900. ( 9a ) POTONIE, Cycadofilices in ENGLER, PRANTL

i.4, 1901. SCOTT, Origin of Seed-Bearing Plants, Roy. Inst. of Gr. Britain, 1903.
OLIVER, Older Gymnosperms, Annals of Bot. xvii. 1903. OLIVER and SCOTT, Phil.
Trans. Roy. Soc. Lond. ser. B. cxcvii. 1904. M. C. STOPES, Flora, 1904. M. BENSON,
Telangium, Ann. Bot. xviii. 1904. C. GRAND_EURY, Pecopteris, Comptes rendus,
140. E. A. NEWELL ARBER, Lagenostoma, Proc. Roy. Soc., 1905, 76. B.
712 BOTANY

(
10
)
and 4
Literature cited under 3 also, PAYER, Traite d'organogenie de la
:

fleur, GOEBEL, Vgl. Entwicklungsgeschichte in SCHENK, Handbuch d. Bot.

1857.
iii. 1, Breslau, 1884. F. NOLL, Normale Stellung zygomorpher Bliiten, etc. 1885,

1887, Arbeit bot. Inst. Wiirzburg, iii. H. VOECHTING, Zygomorphy, Pringsh.

Jahrb. xvii. 1886. VOECHTING, Einfluss des Lichtes, etc., ibid. xxv. 1893.
GOEBEL, Gefiillte. Bliiten, Jahrb. f. wiss. Bot. xvii. 1886. K. SCHUMANN, Bliiten-
morpholog. Studien, Pringsh. Jahrb. xx. 1889. WARMING, Ramification des
Phanerog., Vidensk. Selsk. Skr. ser. v. x. 1872. GOEBEL, Special Morphology, 1882.
J. SCHNIEWIND-THIES, Septalnektarien, 1897. H. MULLER, Befruchtung der Blumen
durch Insekten, 1873. P. KNUTH, Bliitenbiologie, 1 and 2, 1899. C. DETTO,

Ophrys, Flora, xciv. 287, 1905. (") VESQUE, Sac euibryonnaire, Ann. sc. nat. 6 se>.
vi. 1878. STRASBURGER, Befruchtung und Zellteilung, 1878. STRASBURGER,
"
Befruchtungsvorgang b. d. Phanerogamen, 1884. STRASBURGER, Doppelte
Befruchtung," Bot. Ztg. ii. Abt. 1900. STRASBURGER, Befruchtung, Bot. Ztg.
ii. Abt. 1901. L. GUIGNARD, Sac embryonnaire, Rev. sc. nat. Montpellier, T. i.
1882. GUIGNARD, Ann. sc. nat. Bot. 6 ser. xii. and xiii. 1882, and 7 se>. xiv. 1891.
GUIGNARD, vol. jubil. soc. d. Biologic, 1899. GUIGNARD, Antherozoides et double
fdcond., Comptes rend., Paris, 1899. GUIGNARD, Tulipes, Ann. sc. nat. 8 se"r. xi.
1900. GUIGNARD, Maize, Najas major Ranunculaceeae, Journ. de Botan. xv. 1901.
;

TREUB, Loranthace"es, Ann. d. Buitenzorg, ii. and iii. 1883 and 1885. TREUB,
Casuarina, Ann. d. Buitenzorg, x. 1891. TREUB, Balanophora elongata, Ann. d.
Buitenzorg, xv. 1898. S. NAWASCHIN, Birch, Mem. acad. imp. St - Petersbourg,
vii. ser. 42, No. 12, 1894. NAWASCHIN, Elm, Bull. acad. imp. St- Petersbourg,
1898. NAWASCHIN, Lilium Martagon and Fritillaria tenella, Bull. acad. imp.
St- Petersbourg, 1898. NAWASCHIN, Corylus Avellana, Bull. 1899. NAWASCHIN,
Dicotyledones, Ber. d. deutsch. bot. Ges. 1900. WESTERMAIER, Antipodal cells,
Ber. d. deutsch. bot. Ges. 1898. BENSON, Amentiferae, Linn. Soc. London, 1895.
LOTSY, Rhopalocnemis phalloides, Ann. d. Buitenzorg, 2 ser. ii. 1900. G. KARSTEN,
Juglandaceae, Flora, 1902. F. E. LLOYD, Rubiaceae, Mem. of the Torrey Bot.

Club, viii. 1, 1899, and 2, 1902. DOUGLAS H. CAMPBELL, Peperomia, Ber. d. deutsch.
bot. Ges. 1899. DUNCAN S. JOHNSON, Peperomia, Bot. Gaz. xxx. 1900. J.
HAN STEIN, Entwicklung des Keimes, Bot. Abhandlung, i. 1, 1870, Bonn. TH. C.
FRYE, Casuarina, Bot. Gaz. Aug. 1903. JUEL, Ca.suarina, Flora, 1903. P. K.

LOETSCHER, Antipodal Cells, Flora, xciv. 213, 1905. E. STRASBURGER, Drimys,

Flora, Ergzbd. 1905. CH. H. SHATTUCK, Ulmus americana, Bot. Gaz. xl. 1905.
lla
A. HABERMANN, Synergidae, Diss., Bonn, 1905. ( ) Sv. MURBECK, Alchemilla,
Lunds Univ. Arssk. xxxvi. 1901. H. 0. JUEL, Antennaria, K. Svensk. Ak. Handl.
xxxiii. 1900. JUEL, Taraxacum, Bot. Tidsskr. xxv. Kibbh. 1903, and Arkiv f. Bot.
ii. 4, 1904. J. B. OVERTON, Thalictrum, Bot. Gaz. xxxiii. 1902, and Ber. d. deutsch.
Bot. Ges. 1904. C. RAUNKIAER and C. H. OSTENFELD, Hieracium, Bot. Tidsskr. 25,
1903. OSTENFELD, Ber. d. bot. Ges. 1904, 376 and 537. STRASBURGER, Reduction-
division, Sitz.-Ber. Ak. W. Berlin, 18, 1904. STRASBURGER, Apogamy in Eualche-
milla, Jahrb. f. wiss. Bot. xli. 1905. G. BITTER, Parthenogenesis, Bryonia, Abh.
Nat. Ver. Bremen, 1904. H. WINKLER, Parthenogenesis, Ber. deutsch. bot. Ges.
xxii. 573, 1905. M. TREUB, Elatostema, Ann. de Buitenzorg. 2 ser. 5, 1905.
0. H. JUEL, Taraxacum and Cichorieae, K. Svensk. Vet. Ak. Handlingar, xxxix.
1905. Sv. MURBECK, Taraxacum and Hieracium, Bot. Notiser, 1904. 0. ROSEN-
llb
BERG, Hieracium, Ber. d. bot. Ges. vol. xxiv. 1906, 157. ( ) K. GOEBEL,
Cleistogamous flowers, Biolog. Zentralbl. xxiv. 1904. GOEBEL, Viola, Flora, Ergzbd.
12
) J. GARTNER, De fructibus et seminibus plantarum, i. and
1905. ( ii. Stuttgart,

1789-91. A. P. DE CANDOLLE, Pflanzenphysiologie, trans, by ROEI-ER, ii. 212.

 INDEX OF LITERATURE 713

13
F. HILDEBRANDT, Verbreitungsmittel der
Pflanzen, Leipzig, 1873. ( ) Literature

under 3. TH. Knollen- and -Zwiebelgewachse, Berlin, 1850.

IKMISCH, H.
HALLIER, Polyphylet. Ursprung der Sympetalen und Apetalen, 1901, Abh. a. d.
Geb. der Naturw., Naturw. Verein Hamburg, xvi. E. SARGANT, Theory of the
Origin of Monocotyledons, etc., Ann. of Bot. xvii., Jan. 1903. SARGANT, Bot.
Gaz. xxxvii. 1904. K. FRITSCH, Stellungder Monokotylen, Beiblatt, 79. ENGLER'S
Bot. Jahrb. xxxiv. p. 22, 1905. GOEBEL, Streptochaeta, Flora, 1895, Ergbd.
F. KORNICKE, Species and Varieties of Cereals, Handb. d. Getreidebaues by
KORNICKE and WERNER, i. Bonn, 1885. E. BRUNS, Embryo of Grasses, Flora,
1892, Ergbd. SCHLICKUM, Biblioth. Botan. xxxv. 1896. A. NESTLER, Lolium
temulentum, Ber. d. D. bot. Ges. 1898. P. GUERIN, Lolium temulentum, Bot.
Gaz. 28. A. R. WALLACE, Tropical Nature. ALPH. DE CANDOLLE, Ursprung d.
Kulturpflanzen, trans, by E. GOEZE, Leipzig, 1884. J. REINKE, Asparageae,

Pringsh. Jahrb. xxxi. 1897. TH. IRMISCH, Beitr. z. vergl. Morphologic, 1-6,
Halle, 1853-1879. A. F. W. SCHIMPER, Epiphyt. Veget. Amerikas, Bot.
Mitteil. a. d. Tropen. 2, Jena, 1888. C. MEZ, Jahrb. f. wiss. vol. xl. 1904.
CH. DARWIN, Fertilisation of Orchids. G. KARSTEN, Monocotyledonous Trees,
in "Vegetationsbilder," 1. Reihe, 6, Jena, 1903. R. VON WETTSTEIN, Sokotra,
in "Vegetationsbilder," 3. Reihe, Heft 5, Jena, 1905. (
13a
) BUCHENAU in
ENGLER and PRANTL, ii. 1, 228. ) H.
(
13b
GLUCK, Alismaceen, Jena, 1905. ( 14 )
Literature under 3, also : J. LUBBOCK, Contrib. to the Knowl. of Seedlings, vols.
i. and ii. 1892. H. HALLIER, Vorlaufiger Entwurf des natiirl. (phylogenetischen)
Systems der Bliitenpflanzen, Bull. d. 1'herbier Boissier, 2 se"r. iii. 1903. HALLIER,
Umfang, Gliederung und Verwandtschaft der Familie der Hamamelidaceen, 1903.
HALLIER, Phylogeny of Engler's Resales, Parietales, Myrtiflores, etc., Abhandl.
d. Naturw. Vereins, Hamburg, 1903. HALLIER, Daphniphyllum, Tokio Bot. Mag.
xviii. 1904. E. SARGANT, Origin of Monocotyledons, Ann. of Bot. xvii. 1903.
SARGANT, Bot. Gaz. 37, 1904. G. SENN, Hallier's Angiospermensystem, Beih.
Bot. Centralbl. xvii. 1904. K. FRITSCH, Stellung d. Monokotylen, Beibl. 79, in
ENGLER'S Bot. Jahrb. xxxiv. p. 22, 1905. E. STRASBURGER, Drimys, Flora, Ergbd.
15 1Sa
1905, 215. ( ) Cf. 3. ( ) TH. NICOLOFF, Juglande"es, Journ. de Botanique. tt.
xxviii.-xxix. 1904-5. (
16
) H. GRAF zu SOLMS - LAUBACH, Herkunft, Domesti-
kation und Verbreitung d. gew. Feigenbaumes, Abh. d. Kg. Ges. d. W. Gbttingen,
1882. SOLMS -LAUBACH, Geschlechtsdifferenz, etc., Bot. Ztg. 1885. FRITZ
MULLER, Caprificus und Feigenbaum, Kosmos, vi. 1882. G. KING, Ficus, etc.
Linn. Soc. xxiv. G. KING, Species of Ficus, Ann. Roy. Bot. Garden, Calcutta,
1887. A. DE CANDOLLE, Ursprung d. Kulturpflanzen, trans, by GOEZE, 1884. M.
GOLENKIN, Urticaceae and Moraceae, Flora, 1894. 0. WARBURG, Kautschuk-
pflanzen, Berlin, 1900. E. ULE, Kautschukpfl. der Amazonasexped. Engler's
Jahrb. xxxv. 663, 1905. P. REINTGEN, Kautschukpflanzen, Beihefte d. Tropen-
pflanzer. vi. 1905. M. TREUB, Ficus hirta, Ann. de Buitenzorg, 2 ser. A. 3, 1902.
A. F. W. SCHIMPER, Pflanzen und Ameisen, Bot. Mitteil. a. d. Tropen, i. 1888.
17
E. ULE, Ameisenpflanzen, Engler's Jahrb. xxxvii. 335, 1906. ( ) K. GOEBEL,
Verzweigung dorsiventraler Sprosse, Arb. d. Bot. lust, zu Wiirzburg, ii. 3, 1880.
17a des
GOEBEL, Vgl. Entwicklungsgeschichte, 1884. ( ) WIESNER, Rohstoffe
18
Pflanzenreichs, 2nd edition, 1900, ii. 305. ( E. A. BESSEY, Russisch-Turkestan, in
)
18a H. DE
"Vegetationsbilder," 3. Reihe, Heft 2, Jena, 1905. ( ) VRIES, Mutations-
19
theorie, i. 72, and
662, 1901-3.
ii. ( ) M. RACIBORSKI, Cabombeen und
Nymphaeaceen, Flora, 1894. H. L. LYON, Nelumbo, Minnesota Bot. Stud. ii. 1901.
BiisGEN, JENSEN, BUSSE in "Vegetationsbilder," 3. Reihe, Heft 3, Jena, 1905.
20
E. STRASBURGER, Ceratophyllum, Jahrb. f. wiss. Bot. xxxvii. 1902. ( ) E.
 714 BOTANY

JAXCZEWSKI, Anemone, Rev. gen. de Bot. 1892. 0. STAFF, Aconites of India.

Ann. R. Bot. Garden, Calcutta, x. 1905. ( 20a ) E. STRASBURGER, Drimys, Flora,
21
1905, Ergbd. ( ) 0. WAUHIMU;, Mu>katnuss, Leipzig, 1897. J. M. JAXSK, Ann. dc
Buitenzorg, xvi. 1899. ( )
V. HEHX, Kulturpflanzen und Haustiere, 7th edition,
1902. H. SCHEXCK, Mutflmeerhiiume in " Vegetationsbilder," 3. Reihe, Heft 4.
23
( ) H. GRAF zu SOLMS-LAUBACH, Cruciferenstudien, i.-iv. Bot. Ztg. 1900, 1902,
J. M. .M\< KAIU.ANE, Ann.
<24
1903, 1905. HANNIG, Bot. Ztg. 62, i. Abt. 1904. ( )

of Bot. iii. and vii. 1889, 1893. K. GOEBEL, Pflauzeubiolog. Schilderungen, ii.
24 25
1891-93. C *) GOEBEL, -ibul. Wasserpflanzen, 1889. C ) K. GOEBEL, ibid.
) E. DE JANCZEWSKI, Ribes, 1901, Bull. Acad. Cracovie.
<28
i. Sukkulenten, 1889.
(
17
G. TISCHLER, Ribes-Hybriden, Jahrb. f. wiss. Bot. 1906, 42. (-
) K. GOEBEL,
Bot. Ztg. 1882. A. DE CANDOLLE, Ursprung der Kulturpflanzen, trans, by GOEZE,
1884. ('*) TH. BELT, Naturalist in Nicaragua, 1888. H. DE VIUES, Mutations-
theorie, i. ii. 1901-3. G. TISCHLER, Cytisus Adami, Ber. d. deutsch. bot. Ges.
1903. J. REIXKE, Leguininosi-n, i.-vii. Pringsh. Jahrb. xxx. 1896-97. F. NOLL,

PfropfbasUrde, Sitzber. d. niederrhein. Ges. f. Nat. und Heilk. Bonn, 1905. E.

STRASBURGER, Typ. und Allotyp. Kernteilg. Jahrb. f. wiss. Bot. vol. xlii. 1905,
29
62. C ) A. DE CAXDOLLE, Ursprung d. Kulturpflanzen, 1884. V. HEHN, Kultur-
30
pflanzen und Haustiere, 7th edition, 1902. ( ) SCHWEIXFURTH, Erythraea in
S1
"Vegetationsbilder," ii. Reihe 8, Tafel 58, Jena, 1905. ( ) E. WAUMIXI;,
Pollenbild. Phyllome und Caulome in HAXSTEIN, bot. Abh. ii. 2, 1873. R.
BROAVX, Vermischte Schriften, i. 56, Leipzig, 1825. Rubber plants cf. 16. J<>s.
SCHWEIGER, Samenentwickluug der Euphorbiaceen, Flora, xciv. 339, 1905. (**)
33
0. PREUSS, Expedition nach Zentral- und Siidamerika, 1901. ( )
J. WIK..-M i;.

Rohstorte des Pflanzenreichs, 2nd edition, 1900. ( M ) ENGELMANN, Botanical Works

coll. for H. SHAW, edited by W. TRELEASE and ASA GRAY, Cambridge, 1887.

GOEBEL, Pflanzenbiol. Schilderungen, i. Sukkulenten, 1889. G. HEYL, Cacteen,

Archiv d. Pharmacie, 1901, includes further literature. G. KARSTEX and E.
STAHL, Mexikau, Cacteen in "Vegetationsbilder," i. Reihe 8, 1903. E. STAHL,
Mexikan, Xerophyten, ibid. ii. Reihe 4, 1904. ( M ) H. DE VRIES, Mutations-
35a
theorie, i. and ii. 1901-3. ( ) A. K. SCHIXDLER, Halorrhagaceen, Beibl. 77.
Engler's Jahrb. xxxiv. 3, 1904. (
M ) A. F. W. SCHIMPEK, Indo-malayische Strand-
flora,1891. G. KARSTEX, Mangrove veget, Bibl. bot. xxii. 1891. KARSTEX in
"Vegetationsbilder," 2. Reihe, Heft 2, Jena, 1904. G. HABERLAXDT, Maugrove-
pflanzen,Ann. de Buitenzorg, xii. 1893. JOHS. SCHMIDT in "Vegetationsbilder,"
3. Reihe, Heft 7, 1906. ( ')
rH. GRAF zu SOLMS-LAUBACH, Ban und Entwieklung
parasitischer Phanerogamen, Pringsh. Jahrb. vi. 1868. J. SCOTT, Loranthaceae,
Journ. of Agricult. and Horticult. Soc. of India, 1871, and Auszug resp. Ubers.
(Santalum album) by H. GRAF zr SOLMS-LAUBACH, Bot. Ztg. 1874, No. 9. (*)
Literature under 37, also M. TREUB, Loranthacees, Ann. de Buitenzorg, ii. and
:

iii.1883 and 1885. L. JOST, Mistel. Bot. Ztg. 1888. T. JOHNSON, Arceuthobium
Oxycedri, Ann. of Botany, ii. F. JOHOW, Chilen, Bliiteu, Verb. d. Deutsch. wiss.
Ver. Santiago, iv. 1900. L. KLEIN, Forstbotanik, Lorey's Handb. d. Forstwisseu-
schaft, 2nd edition, 1903. K. REICHE, Phrygilanthus aphyllus, Flora, 1904. (*)
H. GRAF zu SOLMS-LAUBACH, Rafflesiaceeu uud Hydnoraceen, Bot. Ztg. 1874.
SOLMS-LAUBACH, Brugmansia Zippelii und Aristolochia Clematitis, Bot. Ztg. 1876.
SOLMS-LAUBACH, Philostyles Hausknechtii, Bot. Ztg. 1874. SOLMS-LAUBACH,
Rafflesia, Ann. de Buitenzorg, ix. 1889. SOLMS-LAUBACH, RafHesia uud Brug-
mansia, Ann. de Buitenzorg, suppl. ii. 1897. E. HEINRICHER, Rafflesiaceae,
Denkschr. Akad. Wien. math, naturw. Kl. Ixxviii. 1905. (*) M. TKEUB, Balanophora
elongata, Ann. de Buitenzorg, xv. 1898. P. LOTSY, Balanophora globosa, Ann. de
 INDEX OF LITERATURE 715

Buitenzorg, xvi. 1899. LOTSY, Rhopalocnemis phalloides, Ann. de Buitenzorg,

41
xvii. 1900. ( ) A. ARTOPOEUS, Ericaceen, Flora, 1903. P. GRABNER, Heide
4
Norddeutschlands, Vegetation der Erde, v. 1901. ( -) F. HILDEBRANDT, Monogr.
Cyclamen, 1898. K. GOEBEL, Regeneration, 1902, Biolog. Centralbl. H. WINKLER,
43
Regeneration, Ber. d. deutsch. hot. Ges. 1902. ( ) A. DE CANDOLLE, Ursprung
der Kulturpflanzen, trans, by GOEZE, 1884. V. HEIX, Kulturpflanzen und
Haustiere, 7th edition 1902. H. SCHENCK, Mittelmeerbaume, Jena, 1905, in
" "
Vegetationsbilder, 3. Reihe, Heft 4. P. KNUTH, Handbuch der Blutenbiologie,
ii. 1899. (**) R. v. WETTSTEIN, Saison-Dimorphismus, Ber. d. deutsch. bot. Ges.
xiii. 303. v. WETTSTEIN, Deszendenztheoret. Unters. i., Denkschrift d. k. Akad.
d. W. Wien, 1900. (
44a
E. GILG, Strophanthus, Tropenpflanzer, 1902, 11.
) E.
45
GILG, H. THOMS. H. SCHEDEL, Strophanthus, Ber. d. Pharmac. Ges. 1904. ( )

Kautschukpflanzen, cp. 16. P. PREUSS, Expedition nach Zentral- und Siida-

merika, Berlin, 1901. Kolonial-Wirtschaftl. Komitee, Kunene-Zambesi-Expedition,
herausgeg. von 0. WARBURG, Kolon. Wirtsch. Kom. 1903. (
M ) M. TREUB,
Dischidia RafHesiana, Ann. de Buitenzorg, iii. 1883. 47
( ) G. J. PEIRCE, Haustoria,

Ann. of Botany, 1893. PEIRCE, Cuscuta, Ann. of Botany, 1894. C

18
) F. MUTH,
Symphytum officinale, Flora, Ergbd. 1902. K. GOEBEL, Boragoid. Flora, Ergbd.
1902. () KOORDERS, Tectona grandis, Natuurk. Tydskr. v. Ned. Ind. 1893.
) A. F. W.
50
C ) M. TREUB, Avicennia officinalis, Ann. de Buitenzorg, iii. 1883. 51
(

SCHIMPER, Pflanzen und Ameisen, Bot. Mitteil. a. d. Tropen, i. 1888. K ) H. (

MULLER, Befruchtung d. Blumen durch Insekten, 1873. M E. HEINRICHER,

( )

Lathraea, Ber. d. k. Akad. d. W. Wien, 1892, and Ber. d. deutsch. bot. Ges. 1893.
HEINRICHER, Halbschmarotzer, i.-iv. Jahrb. f. wiss. Bot. 1897, 1898, 1901, 1902.
R. v. WETTSTEIN, Euphrasia, Ost. bot. Ztschr. 1894, 1895. v. WETTSTEIN,
Monogr. Euphrasia, 1896. STERNECK, Monogr. Alectorolophus, 1901. J. HOFF-
MANN, Odontites, Ost. bot. Ztschr. 1897. K. GOEBEL, Tozzia und Lathraea, Flora,
1897. G. HABERLANUT, Hydathoden, Jahrb. f. wiss. Bot. 1897. 54
( ) HIELSCHER,
M K.
Streptocarpus, Cohn's Beitrage, iii. 1878. ) GOEBEL, Morphol. und biolog.
(

Studien, v. Ann. de Buitenzorg, ix. GOEBEL, Pflanzenbiol. Schilderungen, ii. 116 :

Lentibularieae, 1891.
x
GOEBEL, Flora, 1904, 98. ( ) H. MULLER, Blumen und
S7
Insekten. P. KNUTH, Blutenbiologie. ( ) M. TREUB, Plantes grimpantes, Ann.
de Buitenzorg, iii. 1883. ( M ) M. TREUB, Myrmecodia echinata, Ann. de Buitenzorg,
iii. 1883, and vii. 1888. K. SCHUMANN, Ameisenpflanzen, Jahrb. f. wiss. Bot. xix.
59
1888. ( ) K. GOEBEL, Campanula rotundifolia, Flora, 1896, and Flora, Ergbd. 1905.

HERM. MULLER, Blumen und Insekten. P. KNUTH, Blutenbiologie. R. SCHULTZ,

60
Monogr. d. G. Phyteuma, 1904. C ) F. NOLL, Cucurbitaceen, Landw. Jahrb. xxx.
Ergbd. P. 1901. F. NOLL, Parthenocarpie, Sitzber. d. niederrh. Ges. f. Natur-
und Heilk., Bonn, 190'2. 0. MULLER, Cucurbitaceae, Cohn's Beitrage, vol. iv.
1887. G. BITTER, Bryonia, Abh. Nat Ver. Bremen, 1904. (
61
) M. v. UEXKULL-

GYLLENBAND, Phylogenie d. Bliitenformen und d. Geschlechtsvert. b. d. Kompo-

siten, Bibl. bot. Iii. 1901. A. TSCHIRCH, Kompositen-Antheren, Flora, 1904, Ii.
m ) E. 53
( STAHL, Kompasspflanzen, 1883. C ) C. v. NAGELI, Hieracien, Bot. Mitt,
iii. 1866.
NAGELI, Piloselloiden, ibid. 1867. NAGELI, Piloselliformia, ibid. 1867.
NAGELI, Abstammungslehre, 1884. A. PETER, Hieracium, Engler's Jahrb. v.
C. RAUNKIAER and C. H. OSTENFELD, Bot. Tidsskr. xxv. 1903. OSTENFELD, Ber.
d. D. Bot. Ges. 1904, 376 and 537. 0. ROSENBERG, Ber. d. D. Bot. Ges. xxiv.
w
1906, 157. ( ) W. HEEHING, Baccharis (Diss. Kiel) 1899.
M ) A. F. W. SCHIMPER,
(

Plant-Geography, (^a) 0. H. JUEL, [Antennaria, Kg. Sv. Vet. Ak. Handl. xxx.
1900. ( m ) K. GOEBEL, Pflanzenbiolog. Schilderungen, ii. 1891. ( 67 ) The literature
cited under 9 and 9a.
 SYSTEMATIC INDEX
OF THE

OFFICIAL AND POISONOUS PLANTS

Q Official in Great Britain
+ Poisonous
# Official and poisonous
* before the page indicates a figure.

Triticum sativum, 530

Oryza sativa, *529, 530
aOt Claviceps purpurea, *395, 397 Saccharum officinarum, 530
+ Boletus satanas, 410, *411
+ Amanita muscaria, *412
+ Amanita bulbosa, 412, *413 Spadiciflorae
+ Bussula emetica, 413
+ Lactarius torminosus, 413 + Arum maculatum, *535, 536
+ Scleroderma vulgare, *414 + 'alia palustris, 536
(

Pteridophyta Liliiflorae

Aspidium filix mas, 445, *447, *448,

450 * Colchicum autumnale, *537, *538, 543
# Schoenocaulou (Sabadilla) oilicinalis,
+ Equisetum, *456, 458 543
Q Aloe, species of, *540, 543
Gymnospermae Q Urginea scilla, 543
Smilax ornata, etc., 543
+ Taxus baccata, *490, *491, 498. + Paris quadrifolia, *542, 543
Juniperus communis, *492, 494, 498 + Veratrum album, 541, 543
+ Juniperus sabiua, *493, 494, 498 + Convallaria majalis, *33, 543
Juuiperus oxycedrus, 498 Crocus sativus, 543, *544
Picea excelsa, *495, 496, 498
Abies balsamea, 498
Pinus s}'lvestris, *497,
498 Scitamineae
Pinus palustris,498
Pinus taeda, 498 Zingiber officiuale, 546, *547
Elettaria cardaiuoinuni, 546, 547
Pinus pumilio, 498

Glumiflorae Piperinae
+ Lolium temulentuni, *529, 530 Piper uigruni, *553
+ Lolium remotum, 530 Piper cubeba, *553, 554
717
718 BOTANY

Saliciflorae Rosiflorae

Salix and Populus, species of, *555, *556 Rosa gallica, 598
Rosa damascena, 599
Prunus amygdalus, 598, 599
Querciflorae Prunus serotina, 599
# Prunus laurocerasus, 598, 599
Quercus infeetoria, 653
Hagenia abyssinica, *596, 597, 599
# Quillaja Saponaria, *595, 599
Urticinae

Ficus carica, *565, 566

Leguminosae
Cannabis sativa, 566
Huruulus lupulus, *565, 566 Acacia Senegal, *599, 602
Cassia angustifolia, *601, 602, 605
Cassia acutifolia, 605
Centrospermae Cassia fistula, 605

*568 Copaifera Langsdorfii, *603, 605

Rheum, species of, 567,
Tamariudus indica, *602, *603, 605
+ Agrostemma Githago, *570, 571
+ Haematoxylon campechianum, 605
Saponaria officinalis, *f>71
Krameria triandra, *604, 605
+ Cytisus laburnum, *606, 607, 609
Polycarpicae + Cytisus, other species of, 609
+ Corouilla varia, *609
+ Ranunculus sceleratus, *574, *575 + Wistaria sinensis, 609
+ Ranunculus arvensis and other species, Astragalus gummifer, *607, 609
*574 Glycyrrhiza glabra, *608, 609
+ Caltha palustris, 574, *577 Spartium scoparium, 609
+ Anemone pulsatilla, 574, *576 # Physostigma venosum, 609
+ Anemone nemorosa, *573, 574 Andira araroba, 609
+ Clematis, species of, 574 Pterocarpus santalinus, 609
+ Adonis autumnalis, *573 Pterocarpus marsupium, 609
+ Helleborus, species of, 574, *577 Myroxylou toluifera, 609
# Aconitum Napellus, 577, *578 Myroxylon Pereirae, *604, 605, 606
+ Aconitum lycoctonum. and otherspecies, 609
577, "579
Hydrastis canadensis, 579, *580
& Delphinium staphisagria, 578 Gruinales
Cimicifuga racemosa, 579
Illicium anisatum, 579 Linum usitatissimum, *611
+ Illicium religiosum, 579 Erythroxylou Coca, *611
.Myristica fragrans, 580, *581
Guiacum officinale, 611
Podophyllum peltatum, 580, *581 * Citrus vulgaris, *612
Jatrorrhiza palmata, 580, *582 Citrus aurantium, var. Bigaradia, 614
Cinnaiuomum Camphora, 583, 585 Citrus medica, var. limonum, 614
Cinnaiuomum zeylanicum, 583 Barosma betulina, 614
Sassafras officinale, *583 Cusparia febrifuga, 614
Pilocarpus jaborandi, 614
Picrasma excelsa, 614
Rhoeadinae Quassia amara, *613, 614
Balsamodendron myrrha, 614
# Papayer somniferum, 587 Polygala senega, *614, 615
Papaver Rhoeas, *586, 587
Sinapis alba, 591
Brassica nigra, *589, 591
Cochlearia armoracia, 591 Tricoccae

+ Mercurialis annua, *615

Saxifraginae + Euphorbia, species of, *616, *617

Croton Eleuteria, 618
Liquidambar orientalis, 593 Croton tiglium, 618
Hamamelis virginiana, 593 Q Ricinus communis, 617, *618, *619
 SYSTEMATIC INDEX OF OFFICIAL AND POISONOUS PLANTS 719

Sapindinae Ericinae

+ Buxus sempervirens, 619 Arctostaphylos Uva ursi, *646

+ Rhus toxicodendron, 619 + Rhododendron, 646
+ Euonymus europaeus, *620 + Azalea, 646
+ Ledum, 646
Frangulinae
Diospyrinae
Rhamnus purshianus, 623
Vitis vinifera, 623 + Styrax Benzoin, 646

Columniferae Primulinae
Theobroma cacao, 624, *625, 626 + Cyclamen europaeum, *648
Q Gossypium, species of, *627 + Anagallis arvensis, *648
+ Primula obconica, 648
+ Primula sinensis, 649
Cistiflorae

# Garcinia Hanburyi, 628

Contortae

Olea europaea, 649, *650, *651

Thymelaeinae
# Strychnos nux-vomica, 649, *652
Daphne Mezereum, *631 Gelsemium nitidum, 649
# Daphne Laureola, 631 Gentiana lutea, and other species, 651,
^c Daphne Gnidium, 631 *652
Swertia chirata, 651
4- Menyanthes trifoliata, 651
Myrtiflorae Strophanthus kombe, 651
Strophanthus hispidus, 651, *653
Eugenia caryophyllata, 633, *634 4- Nerium Oleander, 651, *654
Q Pimenta officinalis, 634
4- Vincetoxicum oflicinale, 654, *655
Melaleuca leucadendron, 634
Hemidesmus indicus, 656
Eucalyptus globulus, 632, 634
Q Punica granatum, *635
Tubiflorae
Umbelliflorae Exogonium purga, *656, 657
Convolvulus Scammonia, 657
4- Hedera helix, 636,*637 Lavandula vera, *658, 660
jfc Couium maculatum, *642
Ferula foetida, 642 Galeopsis ochroleuca, *658, 660
Rosmarinus officinalis, 660
Ferula galbaniflua, 642
Mentha 660
piperita,
Dorema ammoniacum, 643
Mentha viridis, 660
Pimpinella anisum, *637, 643 Mentha arvensis, 660
Coriaudrum sativum, *637, 643
Q Foeniculum capillaceum, 643
Q Thymus vulgaris, 660
Monarda punctata, 660
Carum carvi, *638, 643
Q Anethum (Peucedanum) graveolens,643
Ferula sumbul, 643 Personatae
+ Cicuta virosa, *640
4- Sium latifolium, *639, 641 + Nicotiana tabacum, 663, *665, *666
+ Oenanthe fistulosa, *638, 641 4- Lycopersicum esculentum, 661
4- Aethusa cynapium, *644, 641 4- Solanum dulcamara, 661, *663
4- Berula angustifolia, 641 4- Solanum tuberosum, 661
+ Solanum nigrum, 661
# Hyoscyamus niger, 663, *667
Hysterophyta A Datura stramonium, 661, 663, *664
# Atropa Belladonna, 661, *662, 663
Santalum album, 644
Capsicum minimum, 663
4- Viscum album, *644 # Digitalis purpurea, 666, *668, *669
720 BOTANY

Ecballium elaterium, *678

Rubiinae
+ Bryouia dioica, *678
Cinchona succirubra, *671, *672, 673
Uragoga Ipecacuanha, 673, *674
Ourouparia gambir, 673
Sambucus nigra, 675, *676 Aggregatae
Valeriana otticinalis, 675, *676
Auacyclus Pyrethrum, 688
Q Artemisia maritima, var. Stech -

manniana, 688
Campanulinae Anthemis nobilis, 688
Lobelia inflata, *677 Taraxacum officiuale, *683, *684, 688
Citrullus Colocynthis, 678, '679 Arnica montana, *681, *687, 688
 INDEX
(Asterisks denote Illustrations)

Abies, *494, 496 Aeschynanthus, 666

Abietoideae, 494 ^Esculin, 80, 226
Absciss layers, 150 -Esculus, *508, 620; bud -scales, 37;
Absorption, 193 colleters, 109
Absorptive powers of soil, 199 ^Estivation, 38
Acacia, *599, *600 ; gum, 71, 98 Aethusa, *641
phyllode, 45, 163, *210 seedling, *162
; ;
African Oil palm, see Elaeis
spliaerocephala (ant-plant), *235 Agaricineae, 412
Acaciae Gummi, 602 Agathis, 496
Acanthaceae, 668 Agave, 213, 543
Acanthorhiza, root-thorns, 49 Aggeratttm, 682
Acanthus, 668 Aggregatae, 679
Accessory shoots, 20 Agrimonia, 597
Acer, 620, *621 leaf, *32
; ; striatum, Agrostemma, *570, 571
epidermis, 146 Agrostis, 530
Aceraceae, 620 Aira, 530
Acetabularia, *360 Aitionomic movements, 272
Acetic acid, 80 Ajtiga, 659
Achene; *520 Albugo, 382, *383
Achillea, *505, 688 Albumen crystals, 77
Achimenes, 666 Albuminous substances, formation of, 221
Achlya, *381 Alburnum, 133
Achnanthes, 350 Alchemilla, *506, *594, 597 apogamy, ;

Acid substances excreted from roots, 200 93, 517 exudation of water from, 20&
;

Aconiti radix, 578 Alcoholase, 223

Aconitine, 226 Aldrovanda, 591
Aconttum, *505, *573, 577, *578, *579 ; Alectorolophus, 666
exotropism, *282 Aleurone, 76
Acorus, *502, *534, 536 ethereal oil, 79
; ; Algae, 329 assimilation in, 217
; form, ;

root, 121, *114 13 fossil, 465 selective power, 194

; ;

Acrocomia, fruit, *322 Alisma, *518, 523

Actaea, 577 Alismaceae, 523
Actinomorphic flowers, 507 symmetry, ; Alkaloids, 80, 226
15 Alkanet, Anchusa
Adiantnin >'ifi/cn-i>i-//iii, budding, 31 Allium, 541 buds, 20
; ; mucilage, 79 ;

Adonis, *573 roots, *119, 121

Adventitious embryos, 302 ; formations, Allochlorophyll, 62
248 ; roots, 47*; shoots, 20 Allotypic division, 86
jEcidium, 404, *405 Ala us, *557 ;
root tubercles, 232
Aerating roots, 49 Aloe, 213, *540, 543 ; epidermis, *103 ;
Aerial roots, 46 thickening, 144
Aerobionts, 242 Aloes barbadensis, 543 ; socotrina, 543
Aerotropism, 286 Alopecitrus, 530

721 3 A
722 BOTANY

Alpine plants, 213 .1 iiyraecum, 48

Alpine Rose, Rhododendron 631
Aiihtdiiiiitiiii,

Alpinia, 547 Animals, defence against, 116

Alsineae, 571 Anise, Illicit' m unixatiim, I

Alsophila, *446, *448 Anisi stellati fructus, 579

Alternation of Generations, 164, 422; in Anixitropy, 273
Phanerogams, 476 in Rhodophyceae,
;
Annual rings, 131, *132
376 in Thallopliyta, 331
;
scheme ;
.1 a a iilu riii, 466

of, 477 Aunulus, of fern, 448 of musci, 436 ;

Althaea, *504, *626 .1 /mint, 579

Aluminium in plants, 187, 192 Anonaceae, 579

Amanita, *412, 413 Ant-plants, 235
Amarautaceae, anomalous thickening, Anteunaria, 686 apogamy, 93 ;

*143 Anthemideae, 688

Aniarantus, vascular bundles, 124 Anthemidis flores, 688
Aniaryllidaceae, 5i:J Anflif.mix, 688
Amber, 98 Anther, *476, 502
Amber fir, Picea succinifera Antheridia, *371, *372, 375, 421. 4'24, *449
Amicia, movements, *294 Anthoceros, *429
Amides, 80, 222 A nthocerotaccae, 428 stomata. l.vi ;

Amitotic division, 87 Anthocyanin, 80

Ammoniacum, 643 Aitthoxanthum, 530
Amoebae, symbiosis with algae, 235 Anthriscus, 641
Amoeboid movements, *265 Anthurimn, 535
Ampelopsis, *27, *622, 623 ; tendril, *27, Aitthyllis, 607
44, *289 Anticlinal walls, 156
Amplexicaul leaves, 33 Antimony 187
in plants,

Amygdala amara, 599 ; dulcis, 599 Antipodal 514

cells,

Amygdalin, 80, 226 Antirrhineae, 664

Amyloerythrin, 76 Antirrhinum, *519, 665
Amylopectin, 75 Apical cells, *155
Amylum, 530 Apium, 641
Anabaena, 339, 452 Apocarpous gynaeceum, 503
Anacardiaceae, 619 Apocynaceae, 651 latex cells. 7'2
;

Anacyclus, 688 Apogamy, 93, 303, 517

Anaerobic respiration, 241 Apophysis, 436
Anaerobionts, 242 Apothecium, *392, 393, *418, 419
Anagallis, *519, *648 Apposition, 254
Analogous parts, 9 Aqua aurantii floris, 614 rosae, 599 ;

. I 546
Hitiiassa, Aquifoliaceae, 619
Anaphase, 84 Aquttegia, *573, 577
Anaptychia, *420 Araceae, 534 climbing roots, 48 epiphy-
; ;

Anatomy, 52 tic, 213 ;

heat produced by respiration,
Auatropous ovules, *472 244 pollination, 308 velamen, 109
; ;

Anchusa, 658 Arachis, 609 hypogaea, 257, 285

;

.1 ml Ira, 609
Aralia, cauline bundles, 126
Andreaea, *434, 436 Araliaceae, 636
Andreaeaceae, 436 Araroba, 609
Audroecium, 502 .\/-niii-ii-/-iti, 496 wood, 132
;

Andromeda, 646 Arbor Vitae, Thuja occidental i*

Andropogon, 529 Arbuteae, 646
Aneimia, *448 Archaeomlamites, 466
Anemone, *573, 574, 575 ;
adventitious Archangelica, 641
shoots, 20 Archegoniatae, 421
Anemophilous plants, 306 Archegonium, 421, *422, 424, *450
Anethi fructus, 643 Archeplasm, 59
Anethum, 641, 643 Archetypes, 3
Aneura, apical cell, 155 Arctic plants, 213
.
mjiopteris, 445
I
Arctostaphylos, *646
Angiospermae, 478, 501 ; fertilisation, 91, Arcyria, *341
*92 fossil, 688
; Areca, 532
 INDEX 723

-iii, 571 A urii-n.lii, central cylinder, 120

Arenija, 534 Auricularieae, 408
Aryemoiie, 586 Autouomic movements, 272
Arillus, *474 Auxanometer, *255, 256
A ristoloctiia, *643, 644; Howi-rs, *312 ; Auxiliary cells, 376
stem, *118, *128 ; vascular bundles, Auxospores, *348
*129 A vena, 528, 530 starch grains, *74
;

Aristolochiaceae, 643 Avicennia, 659 ; respiratory root, *246

Armeria, *505, 649 Axial wood, 143
Armillai-ia, *400, *409 Axile placentation, 504
Armoraciae radix, 591 Axillary shbots, 20 buds, traces ; of, 124
Amii-a, *680, *681, *687, 688 Axis, 22
Arnicae rliizoma, 688 Azolla, 452
Aroideae, internal hairs, lit! Asotobacter, 189, 232, 337
Arrhenatherurn, 530 Azygospores, 384
Arsenic in plants, 187
Artemisia, *685, 688 Baccharis, 686
Artichoke, Cynara Jerusalem, He/in nt/inx
; Bacillus, *333, *334 ; railicicola, 23'2,
tuberosits 337 ; subtilis, *333
Artocarpus, 566 Bacteria, 331 form, 12
; fossil, 466 ; ;

Arum, *535, 536 ett'ect on substratum of, 232 phos- ;

Asafetida, 642 phorescent, 246

AMI /-I/ in, 644 Bacteriastrum, *350
A.M'fiit of water, 203 Bacterioids, 232
Asdepiadaceae, 653 ; latex cells, 72 Bacterium, 335
Baiera, 501
Ascobolus, 393 r,iil<i,tnfin<irn, <i4") apogamy, 303;

Ascolichenes, 418 Balanophoraceae, 645

Ascomycetes, 387, 388 ;
free cell formation Balata, 646
in, 91 Balsaminaceae, 611
ASCIIS, *90, *387 of, *393
; development Balsamodendron 614 ,

Asexual generation of Bryopliyta, 424 Balsamum peruvianum, 609 tolutanum, ;

Asexual reproduction, 298 609

Ash, 187, 193 Bamboo, Sambusii
A sli, l-'/-ti.i'iiinx Bambusa, 530 rate of growth of, 255
;

Asparageae, 538 Banana, Musa

Asparagin, 80, 201, 222 Banyan, Ficus benyaleitsis, Ficiis indica
AX/HI i-agus, 543 root epidermis, 109
; Barberry, Berber ix rii/i/nri*
Aspery'dlus, *371 ; fumigatus, 233 Barium in plants, 187
Asperula, 673, *675 Bark, 148
Aspidistra, respiration in, 240 Barley, Hardeum
Aspiditim, *439, *447, 448, 450 Jiarosma, 614
Atfifi'iiinni, 251 ; leaf runners, 44 A'/W^.v, ; Bartsia, 231, 666
213 vegetative reproduction, *302
; Rasidiobolus, 386
Assimilation, 214-220 specific energy of,
; Basidiobolaceae, 386
220 ; Engelmann's method of demon- Basidiolichenes, 420
strating, 266 I'xixliliiitnycetes, 387, 400
Aster, 686 Basidium, 388, *400
Astereae, 685 Bast, 138
Astragaleae, 607 Batrachospermum, *375, 376
Astragalus, *607, 608, 609 ; gum, 71 Beaked Parsley, Anthriscus
Astrantia, 640 Beech, fagus
Asymmetrical members, 16 ; flowers, 508 Beet, Beta
Atavism, 165 Besrgiatoae, sulphur in, 79
Atmospheric pressure, 204 Begonia, 47, 630 adventitious roots,
; 17 ;

Atriplex, *570 cauline bundles, 126

A/i-njta, 661, *662, 663 Begouiaceae, 630 ; epidermis, 109
Atropine, 226, 663 Belladonnae folia, 663 ; radix, 663
Atropous ovules, *472 Benzoinum, 646
Attraction sphere, 53 Berberidaceae, 580
Aurantii cortex recens, 614 ; siccatus, 614 Herberts, 580 thorns, 45
;
724 BOTANY

Berry, *520 Brijonia, *678

Bertholletia, albumen crystals, 77 ; fat, 78 Bryony, Bryonia,
Berula, 641 BryopfiyUvm, 592; bmls,j21
Beta, 568, *569 ; etiolation, 259 Bryophyta, 421 form, 14 ; ; fossil, 466 ;

Betula, *557, 558 ; bark, 149 structure, 153

Betulaceae, 557 Bryopsis, 251, 361
Betulin, 149 Buchu folia, 614
Bicollateral bundles, 113 Buckwheat, Fagopynnn
Bidats, 687 Bud, 19, *22, *252 metamorphosis of, ;

Bignoniaceae, 666 ;
anomalous thicken- 23 resting, 22
; scales, -J-J. *36, ;

ing, *143, *144 37, 101

Bilateral flowers, 508 ; symmetry, 16 Budding, 251, *252 ; multiplication by,
Biophytum, 610 301
Birch, Betula Bngloss, AV7//V///
Bird Cherry, Prunus arlum Bulb, *24
Bird's Nest, Monotropa Bulbils, *23, *28, 301
Bird's-nest Orchid, Neottia, Bulbochaete, 358, *359
Bisymtnetrical flowers, 508 Bundles, bicollateral, 113; cauline, 126;
Bitter principles, 80, 226 collateral, 113 ; common, 126 con- ;

Blackberry, Rubus centric, 113 course of, 123-126 ;

;

Blackthorn, Prunus spiimsa foliar, 126 open, 114 ;

terminations ;

Blasia, *14, *4'29 of, 114

Bleeding of plants, 201 Bupleurum, 640
Blepharoplasts, 61 Burdock, /./'/'
Boehmeria, 566 Burnet Saxifrage, I'ii/ijiind!<i
Bogbean, Menyanthes Burseraceae, 614
Boletus, 410, *411 tissues, *97 ; Butomus, 524
Borage, Borago Butter-bur, Petasites
Boraginaceae, 657 Buttress roots, *185
Borago, *657, 658 Buxaceae, 619
Bordered pits, 67 -i's, 619
Boron in plants, 187
Bostryx, *512 Cabbage, Brassica
Bosicellia,614 Cabomba, *573
Botrychium, 443, *444 Cactaceae, 73, 630
Botrydium, *352 Cactus, 27
Boudiera, 389 <',n-xnl/i!n!it, wood, 133.
Bovista, 414 Caesalpiniaceae, 602
Bracken, Pteris aquilina Caffeine, 226, 628
Bract, 37, 509 scale, 495
; Calamarieae, 466
Bracteal leaves, 31, 37 Calamogrostis, 530
Bracteole, 509 Calamostachys, 466
Branch systems, *17 Calcicolous plants, 191
Branched pits, 66 Calcium in plants, 187, 188, 190
Brand spores, *401 Calcium carbonate, 71, 133 excretion of, ;

Brassica, *588, *589, 590; adventitious 104

shoots, 20 Calcium oxalate, 71, '73,' 78
Brazil nut, Bertholletia Calendula, 688
Bread-fruit, Arlocarpus Calenduleae, 688
Bristles, 105 Calla, 536
Briza, 530 Callithamnion, *37f>
Bromeliaceae, 544 ; climbing roots, 48 ; Callitrichaceae, 618
epiphytes, 213 Callose, 69
Br/'iHHS, 530 Calluna, 645
Bromine in plants, 187 Callus, 151 -wood, 151
;

Broom, Spartium Calobryaceae, 431

Brown Algae, Phaeophyceae, 365 Calobryum, 431
Brucine, 226 Caltha, 574, *577
Bruquiera, 632 Calumbae radix, 588
e, 436 acrocarpae, 438
; ; pleuro- Calyptra, 15, 46, 436
carpae, 438 stomata, 154; Calyptrogen, 159
 INDEX 725

Calysteyia, 657 Cassia, *601, 602, 605

Calyx, 502 Cassiae pulpa, 605
Cambium, 128, *129, 131, 142 Cassytlia, 583
Cambogia, 628 Costarica, *561, 562
Campanula, *677 ; epidermal cell, *67 ; Castilloa, 565
silica, 71 Casuarinaceae, 555
Campanulaceae, 675 Catalpa, 666
Campauulinae, 675 Catechu, 673
Camphora, 583 Catkin, 509, *511
Camptosorus, leaf- runners, 44 Caulerpa, 360, *361 ; organs, 249 ;

Campylospermae, 641 structure, 152

Campylotropous ovules, *472 Cauline bundles, 126
Canadian Waterweed, Elodea Cecidia, 165
Cananga, 579 Cecidomyia, 166
Canarium, 614 Cecropia, 235
Canna, *507, *548 starch grains, 74
; Cedrus, 498
Cannabinaceae, 566 Celandine, Chelidonium
Cannabis, 566 Celastraceae, 620
Cannabis indica, 566 Celery, Apium
Cannaceae, 547 Cell, *52 budding, 91 ; division, *82,
;

Cantharellus, 412 87 duration of life of, 263

; filament ;

Caoutchouc, 79, 227, 617, 651 surface and masses, 154 forms, 71 ; ;

Capillarity, 204 fusions, 93 ;

internal pressure of, 179 ;

Capillitium, 340
-
multinucleate, 60, 89 ; nucleus, 59 ;

Capitulum, *510 ontogeny of the, 80 ; -plate, 88 ;

Capparidaceae, 591 -sap, 53, 79 ;

size of, 73 ; uninuclear,
Capparis, *690, 591 89 ; -wall, 63, 88
Caprifoliaceae, 674 Cell -wall, growth of, 254 ;
insertion of,
Capsella, *474, *518, *588, 591 ; Heegeri, 157 ; thickening of,*64
306 Cellular plants, 154
Capsici Fructus, 663 Cellulose, 69
Capsicum, 661, 663 Celtis, 563
Capsule, 436, 519 Centaurea, 682 movements, *296
;

Caragana, 609 Centaury, Erythraea

Carbon, 187 absorption, 214-220
;
in ;
Central cylinder, 113, 117, 160
plants, 188, 189 monoxide, 220 ; Centranthus, 675
Carbonic acid gas, source of, 215 Centricae, 349
Cardamine, *588 adventitious shoots, 20
; Centriole, 53, 61, 86
<
!ardamom, 547 Centrosome, 53, 61, 86, 91
Cardinal points, 176 of temperature, 258;
Centrospermae, 566
CardBus, *680, 681, *682 Cephalanthera, 549
Carex, 525, *526 Cephalanthus, 672
Carica, 630 Cephcdotus, 238, 593
Caricaceae, 630 Cerastium, *513, 571
Carnivorous plants, 236 Ceratophyllaceae, 573
Carotin, 63 Ceratophyllum, 49, 197, 573
Carpel, 37, 472 Ceratium, *343
Carpinus, 558, *559 seedling, *161, *198
; Ceratozamia, *482, 485, *486
Carpodinus, 651 Cercis, *601, 605
Carpogonium, 376, *377, 388, *419 Cereus, *630
Carrot, Daucus Carota C'erinthe,658
Carroway, Carum Ceriops, 632
Carui fructus, 643 Ceropegia, latex, 73
Carum, *638, 640, 641, 643 Ceroxylon, wax, 101
Caruncula, 615 Cetraria, *417, *418, 421
Carya, 555 CJiaerophyllum, 641
Caryophyllaceae, 570 Chaetocladium, 385
Caryophyllum, 634 Chalaza, 472
Caryopsis, 520, 527 Chalazogamy, *515, *516
Cascara Sagrada, 623 Chara, *363, *364 parthenogenesis, 93,;

Cascarilla, 618 365 spermatozoid, *92

;
 726 HOT ANY

Characeae, 59 cell - wall,

; 71 ;
divert Cinnamon tree, Qinna/awmwto
nuclear division, 87 Circulation of protoplasm, "^. 2>i7
'

Chasmogamous Mowers, 309 Circumnutation, 27-

Cheiranthus, *588, 590 Oireaea, (>>.
Cheirostrokits, 467
Cfie/iifniiimn, *475, 586 lociiiiutv.m, -WO; :

laticiferous vessels, 94
Chemotactic movements, 266 Cistiflorae, 627
Chemotropic curvatures, 286 r/.v/,/. s., 629
s
Chemotropism, 286 Ottrulhu, t)7. ,'f>7'.'
s
Chenopodiaceae, 567 anomalous thick-; '/V/v/.v, *612, 613, 614 ;
ethereal oil. !

ening of, *143 Cladodes, 25

Ohenopodiwn, 570 Cladonia, *418, *419
Cherry, /Vc/M/.s- <
'

CZodqp&ora, 359, *360 ; cell, *0 ;

cell
Chervil, C/nn'i-ofi/ii/lfitni division, *89 ; <jloi/n-r<it<i, *12
Chini'M- 1'rimrosr, I'riiiinlii nim-nxi.s Cladostephus, apical cells, 155 ; /(// />//-

Chirata, 651 te<?w, *12

Chiropterophilous plants, 30s Cladothrij; *:',3:?, 334
ChlamydomoHdx, *3f>3 ('lasst-s of Phanerogams, 17 s

Chlamydospores, 388 (
327
'lassilication,
Chloramoeba, *351 OfovorM, *410
Chlordla, *356 Clavarieae, 410
Chlorine, in plants, 187,' 191 Clavicepe, *8W, :;'.'/ ;
twaue, *'.'s

Clilnrocnccum, *355 Cleft leaves, :!:'>

Chlorophyceae, 353 (
'leistocarpae, see I'lia-^

Chlorophyll, 61, 62, 217 grains, *93, ;

(
'leistogamous flowers, 309
*268 preparation of solution of, 61
; Cleistogamy, 518
Chloroplasts, *61, 93 Clematis, 574 cell, *64 vascular system,
; ;

Chlorotic, 191 124, *126

Choanephora, 385 Clerodendron, 659
Cholesterin, 78 Climbing plants, 30
Chondrioderma, *55, 3 1 1
Climbing roots, 48, 249
Chondromyces, *337 Closteriinn, *346
Chondrus, *373, 378 Clostridium, 189, *234
Clwrda, *368 Clover, Trifolinui
Choripetalae, 552 Cloves, 634
Chromatin, 57, 59, 81 'Wo/.s-, 682, *683
Chromium in plants, 187 <'l,m'<>, 657
Chromoplasts, 61, *63 Cobalt, reaction, 208 ;
in plants. 1^7
Chromosomes, 81, *82, *83, *85, 92, 164 ;
Cocae 611
folia,
individuality of, S:',
Cocaine, 226, 611
Cliroococcus, 339, 420 Coccone'is, *349
C/i/-i>n/,/iii/ri/e, 357 Cochineal, 631
Chrysanthemum, 688 Cochlearia, *589, 591
Chrysarobinum, 609 Coco-nut, Cocos
Chrysobalaneae, 598 Cocoa tree, Tlieobr<nn<i
Chytridiaceae, 380 Cocos, *531, *532
Cibotium, 450 Codeine, 226
Cichorieae, 682 Coelospermae, 642
Cichorium, *682 Ccenognmi'trs, "7
<
.

Oicuta, *640, 611 Co/ea, 672, *673

Cilia, 265, 332 Coffeae, 672
Cimicifuga, *573, 579 Coffee, Co/ed
Cimicifugae rhizoma, .17 s Cohesion, movi-ments dependent on, 270
Cinchona, *671, *672, 673 CWa, 625
Cinchonae rubrae cortex, ('<!'' Colchicine, 226
Cinchoneae, 671 <'!,!, icum, *537, *.V5S, 543 ; tulici. 26
Cincinnus, *512 Coleochaete, 359
Cinnamomi Cortex, 6*3 Collateral bundk-s. 113
Oinnanwmum, 582, *585, 688 ;
ethereal CoUnna, *419
oil, 79 Collenchyma, 68, 116, 118, 185
 INDEX 727

Colleters, *108 Cork Oak, Quercus sitber

Colletia, 623 cntciatu, spines, 28
; Cormopliytes, 14
Colocasia, exudation of water from, 210 Cormus, 14
Colocynthidis pulpa, 678 Corn Cockle, Agrostemma
Colouring matters, 226 Cornaceae, 635
Colours of flowers, 80 Cormis, *636
Coltsfoot, Tusx!lagf> Corolla, 502
Columella, 433 Coronilla, *609
Column, 549 Cortex, 117, 146, 160
Columniferae, 623 Corydalis, *475, *508, *586, 587
c,,/i'f,-f(, 609 Corylus, *511, 558, *560 adventitious ;

Combretaceae, 'i-'!2 shoots, 20

Comfrey, Symphytitm Coryplm, 262
Commelinaceae, 536 Cosmarium, *345
Commiphnra, 614 Cotton, Oossyjiin ni
Common bundles, 126 Cotton grass, Eriophorum
Companion cells, *95, 112, 137 Cotyledons, 162, 473
Compass plants, 278 Crambe, *588
Complementary cells, 149 Crassula, 592
Compositae, 680 inulin, 80 laticiferous,
; : Crassulaceae, 592 ; epithema, 123
vessels, 94 pappus, 271 ; Crataegiis, *552, 595 ; leaf, 207 ; thorn,
Compound leaves, 33 28
r
Concentric bundles, *ll. >, 113 Orenothrix, 334
Conferva, 351, *352 Crepis, 682
Conidia, 388 Cribraria, 342
Coniferae, 489 medullary rays, 140 red
; ; Crinnm, root epidermis, 109
wood, 186 resin ducts, 98;
*132 ; ; Crocus, 543, 544 tuber, 25 ;

\vliite wood, 186 Croton, 616, 618

Coniferin, 70, 80, 226 Crown Imperial, Fritillaria
Conii, folia, 642 fructus, 642 ; Cruciferae, 587 ; mucilage, 71 ; idioblasts,
Coniine, -j _''> 73
('mi inm, *637, *642
Oryptogama, 328
Coujugatae, 344 Oryptospora, *394
Conjugation, 91, 346 Cubebae fructus, 554
Connate leaves, 33 Cucumber, Cucumis satin'*
Connecting fibres, 87 L' n <-v HI is, 678

Connective, ."in:;
Cucurbita, 678 etiolation, 259
; pollen ;

Consortium, 4l~> grain, *503 rate of growth of, 255

;
:

Constituents of plant substance, 187 seedling, *285 sieve -tubes, *95 ;

;

Constitution water, 196 tactile pits, *65 vascular bundles, ;

Contact stimuli, curvatures induced by, 124

287 Cucurbitaceae, 677 bicollateral bundles,
;

Contortae, 6-J'l 113

Contractile vacuole, 56 Cupressineae, 492
Cupule, 559, *560
Convolvul:reae, 656 C'upuliferae, 559
Cinir<ili-iilii.t, *656, 657 ; adventitious Curvature, 269 autonomic, 272
;
de- ;

20
shoots, pendent on growth, 272 dependent ;

Copaiba, 605 on imbibition, 270 due to variations ;

Copaifera, *603, 605 in light and temperature, 290 ;

Copper in plants, 187 heliotropic, 276 ; paratonic, 272

Cora, *420 Ci'xi-i'ta, 657 nuropaea,
; 228, *229 ;

CoraUorhuu, 4'.', 231, ~>4'.' : rhizome of, haustoria, 249 reduction of leaves.
;

*24 27 root, 46
;

Corchon'x, *'>'!''
Caspar in, 614
Cordaites, 500 Cuspariae cortex, 614
Cordyline, 54:'. thickening, *14f>
:
Cusso, 599
Corethron, 351 Cuticle, 100 regeneration of, 71
;

Coriandri fructus, 643 Cutinisation, 70

Coriandrum, *637, 642, 643 Cutleriaceae, 369
Cork, *52, *147 cambium, 146 ; Cuttings, 251
728 BOTANY

Cyanophyceae, 337 Dicotyledons, 113 central cylinder, 118 ;

;

I'yatheaceae, 449 medullary rays, 139 phloem of, 139 ;

;

Cyathium, *614 secondary thickening of, *1 42 second- ;

Cycadaceae, 484 spennatozoids, 91

; ary wood, 134 ; vascular bundle, 114,
<
'ycadinae, 484 124 ; venation, 35
Cycadofilices, 466, 500 Dichotomous branch system, 17
Cycas, 484, *485, *486 Dichotomy, 18 ; 'false, 18
Cyclamen, *648 ; regeneration, 152 Dictamnus, 612
Cycle, 42 Itli-tifnut'iiKi, 420

C'yclosporeae, 370 Dictyota, *13, *370 ; apical cell, 155

Cydonia, 595 mucilage, 71
; Dictyotaceae, 370
Cylindrocystis, 345 Itin-rillii, 675

Cymose inflorescences, 510 Differentiation, 171

Cynara, 682 Digenetic reproduction, 298
Cynareae, 681 Digestive glands, 211
Cyperaceae, 524 sheath, 36
; silicae, 191 ; Digitaliu, 226
Cyperus, 525 Digitalis, 665, *668, *669 ; folia, '666
Cypress, Cupressus Dill, Anethum
Cypripedium, 549 Dinoflagellata, see Peridineae
Cystocarp, 376 Dioecious plants, 309
Cystoliths, *68 Dionuea, 237, 591 leaf, *238
; ;
move-
Cystopus, Albugo ments, 295
Cytisus, *604, *606, 607, 609, adami, Dioscorea, *544
*317 Dioscoreaceae, 544; thickening, 144;
Cytoplasm, 53, 57 tubers, 49, *50
Diospyrinae, 646
Dactylis, 530 Diospyros, 646
Dahlia, 687 ; inulin, 80 ;
root tubers, Diplocaulescent, 28
*47 Diplococcits, 336
Dandelion, Taraxacum Diploid, 165
Daphne, *631 Diplostemonous androecium, 507
Durlhigtonia, 238, 591 Dipsacaceae, 680
Date palm, Phoenix Dipsacus, 680
Datura, 661, 663, *664 Dipterocarpaceae, 628
Daucus, 641 chromoplasts, *63
;
Direct nuclear division, 87
Decurrent leaves, 33 Dischidia Rajflesiana, 213, 656 ;

Delphinium, *504, 577, 578 Discomycetes, 391

Dentaria, bulbils, *23 Dissemination of seeds, 318
Dermatogen, 157 Divergence of leaves, 41
Desert plants, 211 excretion of water
;
Diversi florae, 68?
by, 211 Divided lea\<

Desiccation, 195 Dodder, Oiis<-/>/n

Desmidiaceae, 345 movements, 266 ; Dorema, 643
Desmodium, movements, 292 Dormant buds, 23
Deutzia, 593 Doronicinn, 688
Development of the shoot, 18 Dorsiventral, flowers, 508 shoots, 42 ; ;

Dextrorse stem climbers, *284 symmetry, 16

Diageotropism, 281 Dor/ttenia, 564
Diakinesis, 84 Double fertilisation, 305
Diandrae, 549 Draba verna, petites especes, 300
liiiinthus, 571 l>,;ifnena, *541, 543,688; raphides, *77 ;

Duipen&ia, *504 thickening, 144, *145, *146

Diatomeae, 347 form, 12 ; silicon, 191 ; ; Drepanium, *512
movements, 266 Itriiii/iK, 552 ; wood, 136

Diastase, 223 Drip tips, 35

Diatropism, 274 Drosera, 236, 591 digestive glands, 107,
;

Dicentra, 587 108, 109 leaf, *238

;

Dichasium, *512, *513 Droseraceae, 591

Dichogamy, 310 Drosophylluni, 591
])ichotomosiphon, 361 Drupe, 520
Dicotylae, 551 x, 597
 INDEX 729

Dryobalanops, 638 Epipogon, 549

Dryopkyttu&i, 689 Epithema, 123
Duckweed, Lemna Equisetaceae, apical cell, 156 ;
central
biii/i-csnaya, 376, *377 cylinder, 118 ; epidermis, 100 ;
sili-

Duramen, 133 con, 71, 191 ; spores, 271

Duration of life, 261, 262 Equisetinae, 455 ; fossil, 466
Equisetum, *455, *456, *457, 458, 466 ;

Ebenaceae, 646 vegetative cone, *156, *157

EcbaUium, *678 Ergot, Olaviceps
Echinocactiis, 630 Erica, 645
Echinodmrus, *523 Ericaceae, 645
Echinops, 681 Ericeae, 645
Ei'lihtm, *657, 658 Ericinae, 645
Ectocarpus, *369 Erigeron, 686
Edelweiss, Leontopodium Eriobotrya, 595
Egg, 91 -apparatus, 514
; Eriophorum, *525
Eir'/ihornia, 196 Erodium, 610 fruit, *27l
;

Elaeagnaceae, 631 Emi.ni, 609

Elaeagnus, 631 root tubercles, 232
; Eryngium, 640
Elaeis, 533 Erysiphe, ascospore formation, *90
Elaphomyces, 397 Erysipheae, 389, 390
Elasticity of plants, 178 Erythraea, 651, *652
Elateriura, 678 Erythroxylaceae, 611
Elder, Sambucus Erythroxylon, *611
Electrotropisin, 286 Escholtzia, 586
Elettaria, *474, 547 Espdetia, 687
/;//-, 524 Ethereal oils, 79, 98, 226
Elongation, phase of, 253 Etiolation, *259
Embryo, 473, 516, *518 ;
of fern, 439 Eucalypti gummi, 634
Embryology, 154 Eucalyptus, 632, 634, 688; heterophylly,35
Embryonic phase of growth, 248 Eucheuma, 378
Embryo sac, 472, *514 ; nuclear division Eudorina, 355
in, 90 Eugenia, 633, *634
Emergences, 50, 105, 107 Euglena, *339
K^imsa, *385, 386 Eumycetes, 386
Enantioblastae, 536 Euonymus, *620 vegetative
; cone, *30
Endocarp, 519 Eupatorieae, 682
Endocarpon, 418 Eupatorium, 682
Endodermis, 117, *120 Euphorbia, 27, *615, *616, *617 ;
succu-
Endogenous, 20 lent stems, *212
*400
Eiiilfiphyllum, Euphorbiaceae, 615 ; caoutchouc, 78 ;

Endosperm, 473, 516 ; nucleus, 516 latex cells, 72

Endothecium, 503 Euphrasia, 231, 666 ; haustoria, 49
Energy, respiration as a source of, 242 Euryale, 573
Enteromorpha, *357 Eusporangiatae, 443
Entire leaves, 33 Everlasting flowers, Hdichrysum
Eutomophilous plants, 307 Evolution, theory of, 1, 175
Entomophthorineae, 385 Exine, 475
Enzymes, 223 Exoasci, 398
E/ilo-iIra, *498, *499 free ;
cell formation, Exoascus, 398
91 Exobasidium, 409
*434
H/iln'inerum, Exocarp, 519
Epiblem, 100 Exodermis, 120
Epidermis, 100, *100, 160 Exogenous, 20
Epigeal germination, 323 Exogonium, *656, 657
Epigynous flowers, *505, 506 Exothecium, 503
Epilolrium, 632 Exotropism, 281, *282
Epinasty, 272 External influences, irritability to, 273 ;

Epipactis, 549 ;
cell division, *88 upon growth, 257
Epiphyllum, 631 Extrorse anthers, 503
Epiphytes, aerial roots, 48 Exudation of water, *209
730 BOTANY

Faba, 609 Foot, 436, 440

rum,
i
*505, *520, 567 ;
water Forget-me-not, ,!///<
iW/.s-

*190
culture, Formic 80
acid,
Fagus, 559, *561 ; leaf, *122 medullary
;
Fossil Angiosperms, 688 Cryptogams, ;

rays, 141 ; splint wood, 133 winter ;

465 Gymnosperms, 500
;

buds, *22 Foxglove, Itiijitiil'i*

150
Fall of the leaves, Fragraria, 597
Fascicular cambium, 129 Frangulinae, 621
Fats, 78 Fraxinus, 649, *650
Fennel, Foenirn/niii Free, cell formation, 90 ; central placeuta-
Fermentatiou, 233, 243 tion, 504 ; leaves, 38 ; nuclear division,
Ferments, 73, 79, -'2:'. 89
Ferns, Filii-innf, adventitious shoots, 20 ; Freycinetia, 530
apical cell, 156 apogamy, 303 ; ; FritiUuria, 543 leaves, 40 ;

apospory, 304 climbing roots, 48 ; ; Fructus Anisi, 643

concentric bundles, *115 epiphy- ; Fruit, *519 ; of, 270
dehiscence enl,.u. ;

tic, 213 heterophylly, 35

;
leaf- ;
ment 127 epidermis, 109
of, ;

runners, 43 petiole, 123 phosphor-

; ; Frulloiiiii, *430 water-sacs, 213;

escence, 247 root apex, *158 scale ; ; Fucaceae, 367, 371

hairs, 107 stomata, 104 Fuchsia, 632
'
;
vessels, ;

96 Fucosan, 368
Fertilisation, 164, 304, 305, *515, *516, Fucus, '367, *371, *372 centriole, 61. ;

*517 *86, '87 nucleus, *53, '86, '87

;

Ferula, 641, 642, 643 Fidigo, 341

Fe*t,i f ,,, 212, *527, 530 Fumaria, *520, 587
Fibre, *72, 81, 136 -tracheides, 72, 135 ; Funiariaceae, 587
Fibro-vascular bundles, 111 Funaria, *61, *91, '423, 438
Fibrous cells, 136, 139 Fundamental tissue system, 100, 115
Fictts, 564, *565 ; -iiulifu, 49 ; pollination, Fungi, cell fusions, 96 effect on sub- ;

308; cystoliths, *68, 71, 109; epi- stratum of, 232; glycogen, 7-
dermis, 109 ; stipules, 37 nutrition of, 234 phosphorescent, ;

Fig, Finis 246 structure, 152 tissues, 97

; ;

Filmjii, 686 Fungus gardens, 235

Filament, 502 Funiculus, 472
Filices, 445 Funkia, '516; adventitious embryos, *303
Filicinae, 443 ; fossil, 466 Furze, Clex
Filix mas, 450
Fir, I'iceu
Flagell", 2f,r, Galbanum, 642
Flagellata, 3:;<< Galegeae, 607
Flax, Litium Galeopsis, *658, 659
Floral, diagram, *40, *508 ; formula, 508: Hull a ni, *520, 673 stipules, 36 ;

leaf structure, 121 ; leaves, 31, 37 ; Galls, 165, 563

shoots, 28 ; symmetry, 507 Gametes, 91, 331
Florideae, see Rhodophyceae . Gametophyte, 164, 439
Flowers, 28, *502 ; colour of, 80 de- ; Oarcinia, 628
velopment in darkness, 259 of ;
Gaseous exchange, mechanism of, 221
Angiospermae, 501 of Gymnospermae, ; Gasteromycetes, 413
480 opening and closing of, 291
; Geaster, *414
Fluctuating variations, 165, 300 Gelsemii radix, 649
Fluorine in plants, 187 Gelsemium, 649
Foeniculi fructus, 643 Gemmae, 23, *426
Foeniculum, *505, *637, 641, 643 Generation, alternation of, 164 diploid, ;

Foliage leaves, 31 ;
structure of, 121, 165 haploid, 165
;

*122 ; metamorphosis of, 43 Generative cells, 476

Foliage shoots, 28 Genetic spiral, 42
Foliar bundles, 126 Genista, 213, 607
Follicle, 519 Genisteae, 607
Fontinalis, 438 Gentiana, 649, 651, *6.v_>
Food of plants, constituents of, 188 Gentianaceae, 649
Fool's parsley, Aethusa Gentianae radix, 651
 INDEX 731

Geotropic, curvature, *280, *2S1 : varia- on, 260 influence of temperature on,
;

tion
movements, 29:> 257 ;
22 ; in thickness,
intercalary,
Geotropism, 278 128, *131, 142; limited, 30; of cell
Geraniaceae, 610 wall, 64 ; mechanical influence on,
Geraniii'ui, *507, *610 260 ; phases of, 248 ; unlimited, 30
Germination, 161, 162, 318, 320, 474 Gruinales, 610
Gesneriaceae, 666 Guarana, 620
Oeum, 597 Guard cells, 103, 206
Gigartina, *374, 378 Guiaci, lignum, 611 ; resina, 611
Ginger, Ziiiyiber, 547 Guiacum, 611
Ginkffo, *481, *488, *4M9 ; spermatozoids, Gum, 70, 98
91 Gummosis, 70
Ginkgoaceae, 488 Gunnera, central cylinder, 120
Ginkgoinae, 488 Gutta percha, 79, 227, 646, 652
Gladiolus, 543 Guttiferae, 628
Glands, 108 Gymnadenia, 549
Glandular hair, *107, 108 Gymnocladus, biids, 20
(it, III: 'ill,,!. *58f> Gymnodinium, *343
Gleba, 413 Gymnogramme, 101
Glechoma, 659 Gymuospermae, 113, 478, 480 ;
fertilisa-

Gleditsehia, 232, 605 ; buds, 20 ; thorns, tion, secondary wood, 134, 142
91 ; ;

*27, 28 central cylinder, 118 fossil, 500 ; ;

Globoid, *76 medullary rays, 139 ; phloem, 138 ;

Gloeocapsa, *11, *338 root apex, 159 ; vascular bundles,

Gloxinia, 666 114, 124
Glucose, 79, 195, 218, 221 Gymnosporangium *405 ,

Glucosides, 80, 225 Gynaeceum, 502

Glume, 526 Gynandrae, 548
Glumiflorae, 524 Gynostemium, 549
Glutamin, .^0 Gyromitra, 394
Glycogen. 7*
Glycyn-hi-M, *608, 609 Haastia, 686
Glycyrrhizae radix, fiOi* Habit of plants, 29
Gnaphaliuw, 686 Hadromal, 70, 112
Gnetaceae, 498 ; vessels, 96 Haematococcus, *354
Gnetinae, 498 Haematoxyli lignum, 605
Gnrfinn., 498, *499, *500 anomalous ; Haematoxylon, 133, 605
thickening, *143 Haemoglobin, 62
Gonidia of Lichens, 416 Hagenia, *596, 597
'"///', *343 Hairs, see Trichomi'-
Gossypiuin, 626, *627 hairs, 105, *106
; Halimeda, 361
Gossypiiim, 627 Halophytes, *192 .

Gracilaria, 378 Halorrhagidaceae, 632

Graft-hybrids, 252, 318 Hamamelidaceae, 593
Grafting, 251, *252 Hamamelidis cortex, 593 folia, 593 ;

Gramineae, *526 epidermis, 100

:
;
root Hancornia, 651
apex, *159; silica, 71, 191; sheath, 35 Haplocaulescent, 28
Granati cortex, &''>:>
Haploid, 165
Grand periods of growth, 254, 256 Haplomitrivm, *430, 431
Granular plasma, 58 Haplostemonous androecium, 507
Grape vine, T///X Harveyella, 378
Graphis, 418 Haustoria, 49, *229
Grass haulm, curvature of, 284, *285 Hazel, Corylus
Gratiola, 665, *667 Heart- wood, 133
Gravity, irritability to, 279; reaction to, 174 Hedera, 636, *637
Ground nut, Ai-n<-h!s lii/i>ogaea Hedychimn, 547
Growing point, 154 Hedysareae, 609
Growth, 247-263 ; by apposition, 64 ; by Heliantheae, 686
intussusception, 64 correlation of,
; Helianthemvm, *628, *629
249 curvatures, 272
; influence of ; Helianthus, *517, *682, 687 ; tuber, 25
light on, 258 influence of moisture
; Helichrysum, 686
732 BOTANY

Heliotactic movement, 277 Hydathodes, 108, 123, 210

Heliotropism, 274, 275 Hydneae, 410
ffeliotropium, 658 Hydnaphytuni, 672
Helleborus, *36, 574, *577 ; epidermis, Hydnum, *410
*102 ; stoina, *206 Hydra, symbiosis with algae, 234
Helobiae, 522 Hydrangea, 593
Helvellaceae, 394 Hydrastine, 579
Hemerucallis, *476 Hydrastis, 579, *580
Hemidesmi radix, 656 Hydrastia rhizoma, 579
Hemidesmus, 656 Jlydrodutri-s, 524
Hemlock, Conium Hydrocharitaceae, 524
Hepaticae, 424, 425 apical ; cell, 155 ; Hydrocotyle, 640
conducting strands, 110 ; elaters, Hydrocyanic acid, 226
271 ; form, 14 Hydrodictyon, 356
Herbs, 29 Hydrogen, in plants, 187, 188, 189
Hercogamy, 311 Hydrolapathum, *13 structure, 1;V2
;

Heriiiaria, 571 Hydrophilous plants, 307

Hesperidin, 80 Hydrophyllaceae, 657
Heterobasidion, 412 Hydropterideae, 450, 466
Heterocontae, 351 Hydrotropism, 286
Heterodynamic hybrids, 313 Hygroscopic movements, 271
Heteroecious Uredineae, 406 Hylocomium, 438
Heterogenesis, 300 Hymenolichenes, 420
Heteromerous Lichens, 416 Hymenomycetes, 408
Heterophylly, 35 Hymenophyllaceae, 449
Heterosporous Pteridophyta, 441 Hyoscyami folia, 663
Heterospory, 471 Hyoscyamus, *474, 663, *667
Heterostyly, 310 Hypecoum, 587
Heterotype division, 86 Hypericum, *628
Hevea, 617 Hyphae, 255, 38" rate of growth of, 255
;

Hibisceae, 626 Hyphaene, 530

Hibiscus, 626 Hypholoma, *400
Hieracium, *682 ; apogamy, 93, 518 ; H A onum, *435, 438
apospory, 518 Hypocotyl, 162
Hilum, 474 Hypoderma, 118, 121
Hippocastanaceae, 620 Hypogeal germination, 323
Hippophiie, 631 Hypogynous, *505, 506
Hippuris, 632 vegetative cone, *157
; Hyponasty, 272
Histology, 52 Hypophysis, 516
Holcus, 530 Hysterophyta, 643
Homodynamic hybrids, 313
Homoiomerous Lichens, 416 Iberis, 591
Homology, 9 ice, formation of, in tissues, 176
Homosporous Pteridophyta, 441 Idioblasts, 73, 116
Homotype division, 86 Ilex, *619
Honeysuckle, Lonicera Illidum, 579 ;
ethereal oil, 79
Hoodia, 656 Imbibition, movements dependent on, 270 ;

Hop, Humultis water, 194

Hordeum, 528 root apex, *159
; Imbricated leaves, 38
Hornbeam, Carpin-us f,,ij>ufie)is, 611 ; bundles, 109 ; *116 ;

Horse-chestnut, Aesculus collenchyma, *67

Horse-tail, Equisetum India-rubber tree, FICHS
Hoya, 656 Indian cress, Tropaeolum
Humboldtia, 235, *236, 605 Indian hemp, Cannabis
Humuhis, *565, 566 glandular ; hair, Indican, 226
107, 108 Ind-igofera, 609
Humus, 199 plants, 231
; Indirect nuclear division, 81
Jfura, fruit, 320 Indusium, 448
Hyacinthus, 541 Inflorescence, 509
Hyaloplasm, 58 Insectivorae, 591
Hybridisation, 313, *315 Insectivorous plants, 43, 227, 236
 INDEX 733

Integuments, 472 Labiatae, 659

Intercalary growth, 257 Labiatiflorae, 682
Intercellular spaces, 97, 98, 244 Laboulbeuiaceae, 388, *398
Interfascicular cambium, 129 Laburnum adami, *317
Internal, development of organs, 248, 260 ; Lachnea, *392
116; morphology, 52 structure,
hairs, ; Lactarius, 412
the phylogeny of, 152 Lactuca, 682 scariola, 277
; ; laticiferous
Internodes, 22 vessels, 94
Intine, 475 Lagenostoma, 509
Intramolecular respiration, 241 Lamina, 31
Intussusception, 254 Laminaria, *366
Introrse anthers, 503 Laminariaceae, 366 sieve -tubes, 110 ; ;

I/nitit, 686 structure, 153

Inuleae, 686 Lamium, *657, 659
Inulin, 80 Landolphia, 651
Iodine, in plants, 187, 373 reaction, *218; Lantana, 659
Ipecacuanha, 673 Lapartea, 566
Ipomfya, 657 Lappa, *681, 682
Iriartec, root hairs, 49 Larch, Larix
Iridaceae, 543 diagram, *40, *41
; Larix, *481, 496 short shoots, 21;

Iris, 543, *545 root, *120

;
Latent buds, 23
Iron, bacteria, 243, 335 in plants, 187,
; Lateral, geotropism, 281 roots, 47 ;

188, 190 Latex, 73, 98, 227 cells, *72 :

Irritability, 4, 174, 273 Lathraea, 665 albumen crystals, 77

; ;

Isatin, 226 exudation of water, 210

Isatis, 591 Lathyrus, 609 stipules, 44 ; tendrils,'44,
;

Isoetaceae, 464 45, *163

Isoetes, *464, *465 desiccation, 195
;
Laticiferous vessels, 94, *96
Isogamous fertilisation, 331 Laudatea, 420
Isosmotic coefficients, 180 Lauraceae, 580
Ivy, Hedera Laurocerasi folia, 599
Laurus, 582, *584 ethereal oil, 79 ;

Jaborandi folia, 614 Lavandula, 658, 660

Jalapa, 657 Lead in plants, 187
Jambosa, 633 Leaf, base, 31, 35 ; blade, 31 ; cushion,
Jasioae, 677 33development of, 30, 31 foliage,
; ;

Jasmiintm, 649 32forms of, 31

; mechanical tissues ;

Jatrorrhiza, 580, *582 of, *184; rigidity of, *185 scars, ;

Juglandaceae, 554 38, 150 sheath, 31, 35

;
stalk, 31 ; ;

Juglandiflorae, 554 symmetry of, *16 tendrils, 44 tip, ; ;

Juglans, *516, *554, 555 31 thorns, 45

; traces, 123 vena- ; ;

Juncaceae, 538 tion, *207

Juneu>.'inaceae, 524 Leaves, arrangement of, 39 autumnal ;

.In in- us, *536, 538 changes, 62 fall of the, ;150 of ;

Juugermanniaceae, 429 mosses, 432 origin of, 160 ; reduc- ;

Juiiijjerus, *492, *493 ;

vascular system, tion of, 213 secondary growth of,
;

123, *124 145 structure of, 121

; succulent, ;

Juvenile form, 163 213

Ledmn, 646
Kalmia, *313 Legume, 519
K'luili'lni seedling, *321 Leguminosae, 599 aluminium, 192 root ; ;

Karyokinesis, 81 tubercles, 232

Kickxia, 651 Lemna, 536 chlorophyll grains, *268
;
;

Kino, 609 root pocket, 46

Kinoplasm, 59, 88 Lemnaceae, 536
Kh'iitM Hrtiri'lata, 213 Lentibulariaceae, 668
Klitiostat, 256 Lenticels, *149, 244
Knautia, 680 Leocarpus, *342
Knots, 144 Leontodon, flower head, *291
Krameria, *604, 605 Leontopodium, 686
Krameriae radix, 605 Lepidium, *588
 734 BOTANY

Lepidodendreae, 466 Lain: in, *529, 530

Lefithutrobug, 467 /.'</'/
/<v/w, 074; liuds, 20; leaves. :;:i

412
/.//iota, Lonicereae, 674
Leptome, 112 Lophvspermvni, *290
Leptomiii, 79 Lorauthaceae, 644
Leptosporangiatae, 445 L&ranthus, 644
l.,/>tothrix, *11, 335 Loteae, 607
/..
Hernia, 336 Lutus, *604, 607
Lettuce, Luctm-n Lovage, Leoittimm
Leucadendron, 212 I.inaria, *588
Ln lii n us, 607
Leucojtim, 543 ; *544 Lnpulin, 108
Leuconostoc, *334 Lupulinum, 566
Leucoplasts, 61, 63 Lnpulus, 566
Li'1-ixtii'nin, till l.ii-.iiln, 538

Lianes, 30 ; vessels, 96, 137, 202 l./li-lmix, 571 i-i.ti-nriii, 101

:

Libriform fibres, 136 Lycopenlon, *414

Lichenes, 415 aluminiuni, ;
192 ; com- Lycopersiciim, 661
position of thallus, 234 Lycopodiaceae, 46, 459 ; dichotomy, 19
Li'-Hiuphora, 348 Lycopodiuae, 458, 466 growing point, ;

Life, conditions of, 172 157 ; root apex, 158

Light, effect in photosynthesis, 216 ;
influ- Lycopodium, *19, *440, *459, *460 ;

ence on growth, 258 ; irritability to, aluminium, 192 ;

central cylinder, 113
274 Lycoi-is, root epidermis. ID'.'

Lignirication, 70 500
LygiiKiili-nilriiii.
Liiniitied cell walls, 69 Lygiuopterideae, 500
Ligule, *34, 526 Lygodium, growth of leaf, 30
Ligulittorae, 682 Lysimachia, 648
l.i<liifitrviii, 649 Lythraceae, 632
Liliaceae, 538 diagram, 40 ; raphides, 78
; Ly thrum, 632
Lilieae, 538
Liliiflorae, 536 secondary growtli of, 127
; Madura, wood, 133
Lilii'ni, *85, *517, 541 bulbils, 23 ; ; Macrocystis, *366
nuclear division, 85 Macrosporangia, 441, *4;j:5, 472
Lily, Liliitm ;
of the Valley, Convallaria Macrospore, 441, *453, 463, I7l!
Lime, Tilia Macrosjwrophyll, 472
Lime scales, 211 Magnesium in plants, 187, 188
651
/.i,iinii,it/tenwm, Magnolia, 579
Liinonis cortex, 614 succus, 614 ; Magnoliaceae, 579
Linaceae, 610
l.iiinria, *511, 665 ; cymbalarm, 257 ; Maize, Zea
heliotropism, 278 Malarophilous i>lants, 308
Linin, 59 Malic acid, 80
I.i.n-Hi, *505, 610, *611 Mnllotiis, 618
Linuin, 611 Maltose, 221
Liquidairibar, 593 Mnli-n, *625, *626 ; Leliotropism, 277 ;

Liriodendron, 579 mucilage, 71 pollen-grain, *503

;

Listera, 549 Malvaceae, 626

Lithium in plants, 187 Malveae, 626
l.ltnrella, 668
Mn miUii fin, 630
Liverworts, Hepaticae, Manganese in plants, 187
Loasaceae, stinging hairs, 106 Mi in'j ifera, 619
Lobed leaves, 33 Mangostcen, t Inn- in in
I.nh,'Ua, *504, *677 Mangrove, roots, 49 seedlings, *321 ;

Lobelia, 677 Manihot, 617

Lobeliaceae, 677 Mannite, 373
Loculicidal dehiscence, 519 Mantle leaves, 35
Locust, Robina pseudacacia Maples,
Lodicules, 526 Maquis, 660
Lmloicea seycliellarum, 178 Mn rait ta, 547
Loiruniaceae, 649 Marantaceae, 547
 INDEX 735

Marattiaceae, 445 pseudo-periderm, 148

; Micellae, 254
M,n;h,u,tia, *421, *422, *426, *427 ; Micrasteruis, *345
air-pore, 152 ; gemmae, 249 Mii-riii-nfi-iia, *11

Maivhantiaceae, 426 Micropyle, 472

Mni-i-iiliiui/i, 660 Microsomes, 58
Mtirsiliii, *451, 466 ; parthenogenesis '.:;, Microsporangia, *441, *453, 47-~>
303 Microspores, 441, *453, 462, 475
.Marsiliaceae, 451 Microsporophyll.s, 47-~>
Mutficariu, *681, *685, 688 Middle lamella, 69, 98
Mufthi'ilti, 590 Midsummer growth, 131
Maximum, intensity of stimulus, 175 ;
M igiionette, Reseda
temperature, 258 Milfoil, Achillea
Mechanical, influence on growth, 260 ; Millet, A ndropogon
rigidity of plants, 116 tissues, 182,
: Millon's reagent, 57
*183 Mimosa,* 599, 600 ; leaf, 34 ; movements,
Mi-i-HiinjLii.i, 286 293, *294, 295
Median plane, 508 Mimosaeae, 599
Medicttgo, 607 Mimusops, 646
.Medlar, Mesp'dus Mineral substances, 195 ; absorption of,
Medulla, 117, 118 195
Medullary rays, 118, *130, *138, '139, Minimum intensity of stimulus, 175 ;

140; sheath, 131 temperature, 258

Medulloseae, 500 Mistletoe, Viscum album
Melnli'i/'-'/, 634 Mitotic nuclear division, 81
Mi'luiii/ii/fiiiii, 231, 666 Mitiitui, conducting strands, 110 ; >tem,
Melanthieae, 538 *153
Melastomaceae, 632 Moisture, influence on growth, 260
Melica, 530 Mdiiiia stem, *183
Mdilolus, 607 Monandrae, 549
Melissa, *660 Moiiurda, 660
Melon, t'ni-Hiiiiti Mdo Monkslipod, 1'" a it uni
. 1

Mftii.tii-'t, 350 .
Monoblepharideae, 379
Members of independent origin, 15, 49 Mniioblepharis, *379
Mendel's laws, 314 Mouochasium, 512
Mriiispermaceae, 580 Monocotylae, 521
Men tlm, 660 Monocotyledons, central cylinder, 118 ;

Menthol, 660 secondary growth, 127, 144 vascular ;

Menyantlifs, 651 bundles, 114, 125 ; venation, 35

Mercurialis, *615, 616 ; epidermis, *102 Monoecious plants, 310
Mercury in plants, 187 Monogenetic reproduction, 298
Meristem, 99 Mouopodial branch system, 17
Merogony, 304 Monopodium, 17
Merulins, 412 Monotropa, 646
Maeatbryanthemvm, 213 Monotropeae saprophytic, 231
Mesocarp, 519 Monstera, 534 leaf, 35 ;

.]/('.wi'<o-/j//x, orientation of chloropla^ts, Monstrosities, 165 causes of, 16b'

;

268 Moraceae, 563 caoutchouc, 79

; ; latex
Mi-sophyll, 121 sheath, 122
; cells, 72
Mesotaeniaceae, 344 Morchella, *393, 394 ascus, *387 ;
'

Mesotaenium, 345 Morphine, 226

Mi'njiiln.f, 595 Morphology, 6, 9, 10
Mestome, 111 Morns, *521, 563
Metabolism, 186, 225 Moss capsule, 424
Metamorphosis, 10 of the bud, 23
;
of ; Mosses, Mitsci
foliage leaves, 43; of primary members, Mousetail, Alyosnrus
15 of roots, 48
; theory of, 15
; Movement, phenomena of, 263-296
Metaphase, 84 Mucilage, 70, 79 ; tubes, 94
Metaplasm, 57 Mucilaginous cell walls, 70 sheath, 321 ;

Metroxylon, 533 Mucor, 383, *384, *385

Metzgeria, 429 ; apex, 155 Mucorineae, 383
Mezerei cortex, 631 Mucunn, anomalous thickening, *143
736 BOTANY

Mulberry tree, Morns Netted veined leaves, 34

Mullein, Verbuscum Nettles, Urtica
Multicellular hairs, 107 Nickel in plants, 187
Multilateral symmetry, 15 Nicotiana, *505, 662, *665, *666
Multiuucleate cells, 60 Nicotine, 226
M ><', 546 ; leaf, 35, 184 Nightshade, Solatium /i/<tn/ni Enchanter's, ;

Musact/at', 546 Cireaea ; Deadly, Atropa

Muscari, 541 buds, 20
; Nitdla, 363 turgidity, *180
; rotation of ;

Muscarine, 226 protoplasm, 59, 267

Mi/fi-i, 426, 431 apical cell, 156
;
con- ; Nitragin, 232
structing strand, 110 stem, *153 ;
Nitrate bacteria, Nitrobacter
Mushroom, I'fid I iota campestris Nitrite bacteria, NitrosmiH'imx
Mustard, Xiim/iis Nitrobacter, *337
Mutations, 3, 300 Nitrogen, 187 ; fixation of, 187, 23'2 : in
Mutisieae, 682 plants, 188, 189
M \cdium, 386 Nitrosomonas, *337, 243
Mycorhiza, 231 Nitzschia, 351
Mynuecophytes, 235 Nodes, 22
Myosotis, 658 ; dorsiventral shoots, 43 Normal shoots, 20
574
)[i/<isurus, Nostoc, *338 ; symbiotic, 429
Myricu, 555 wax, 101
; Nurellus, 472
Myricaeeae, 555 Nuclear, cavities, 83 ; division, 81-87,
MyriophyUum, 632 *82 ; division, allotypic, 86 ; division,
Myristicu, *475, 579, 580, *581 ; fat, 78 direct, 87 ; division, indirect, 81 ;
Myristicaceae, 579 division, heterotype, *85, 86 ; division
Myrmecodia, *237, 672 homotype, 86 ; division, mitotic, 81 ;
Myrobalans, 632 membrane, 60 ; network, 81 plate, ;

Myrouic acid, '226 *82 sap, 60 spindle, 81, *83, *85

; ;

Myroxylun, *604, *605, 606, 609 Nucleoli, 59, 82

Myrrha, 614 Nucleus, *53 albumen, crystals in, 77
;

MyrriphyUum, *284 Nuphar, 572

Myrtaceae, 632,, Nut, *520
Myrtiflorae, 631 Nutations, 272
Myrtle, Myrtus Nutmeg, Myrifitlcfi
Myrtus, 632, *634 Nutrition, 186 special processes of, 227
;

Myxamoeba, 341 Nux Yomica, 649

Myxobacteriaceae, 337 Nyctaginaceae, anomalous thickening,
Myxococcus, 337 *143
Myxomycete, 54, 96, 340 Nyctitropic movements, 293
yymphaea, *475, *572, *573
Naiadaceae, 524 Nymphaeaceae, 572 internal hairs, 116 ;

Aairts, 524
Jfarcissus, 543 Oak, Quercus
Nastic curvatures, 270 Oat, A vena saliva
Nasturtium, adventitious shoots, 20 Obturator, 615
Natural selection, theory of, 2 Ochrea, 37, *567
Stn-iada, *348 Ocimuin, 660
Neckera, 438 Odontites, 666 ; haustoria, 4!

Nectaries, 102, 109, 211, 507 Odontospennitm, 686

395, 397
S<-<-irin, Oedoyonium, 358, *359
Negative, geotropism, 279 ; heliotropism, Oenanthe, *638, 641
275 Oenotliera, *632 lamarckiana, 300
;

Xelumbium, 196, 573 Oidium Tiickeri, 390

Xeuttia, 62, 231, 549 Oil-ducts, 98
Nepenthaceae, 591 Oils, 78, 225
Nepenthes, *43, 591 ; leaf, *238 Olea, 649, *650, *651 ; latent buds, 23
Xepeta, 659 Oleaceae, 649 ; albumen crystals, 77
yephromium, 420 Oleander, Nerium oleander
Nerium, 651, *654 ; stomata, 154 Oleum, cadinum, 498 cajapnti,
;
634 ;

Nervature, see Venation crotonis, 618 ; eucalypti, 634 ; juni-

Nest leaves, 35 peri, 498 ; lavandulae, 660 ;
meuthae
 INDEX 737

660 menthae viridis, 660

piperitae, ; ; Oxalis, 610 movements, 292
; organic ;

olivae,649 pini, 498

; ricini, 618 ; ; acid,80
rosae, 599 rosmarini, 660
; santali, ; Oxydases, 223
644 ; terebinthinae, 498 theo- ; Oxygen, 187 in plants, 188, 189 liberated
; ;

bromatis, 626 in photosynthesis, *219

Olibanum, 614
Olive, Olea Padina, 370
Olpidium, *381 Paeonia, *502, 577 ; flower, *38
Onagraceae, 632 Palaguium, 646, *647
Onoclea, 446 spermatozoid, *92
; Palea, 526
Ononis, 607 Palisade cells, 121, 122; parenchyma,
Ontogeny, 2, 9 ; of the internal structure, *122
154 ; of the cell, 80 ; repeating phy- Palmae, 530 secondary growth of, 127
; ;

logeny, 162 type of vascular system, 125, 126 ;

Oogamotis fertilisation, 331 siliceous bodies, 78 stability, 178 ; ;

Oogonia, 331, *371 wax, 101

Oomycetes, 379 Palmate leaves, 33
Oosphere, 331 Pandanaceae, 530 ; secondary growth of,
Oospore, 331 127
Ophioglossaceae, 443 Pandatius, 530, *533 ; adventitious roots,
Ophioglossum, 443, *444, *445 ;
adven- 49 leaves, 40
;

titious shoots, 47 ; cork, 148 Pandorina, 355

Ophrys, 549 Panicle, *510, *511
Optical apparatus, 102 Panicum, 529
Optimum, intensity of stimulus, 176 ; Pansy, Viola tricolor
temperature, 258 Papaver, *475, *586, 587 ; laticiferous
Opnntia, *26, 630, 631 cladodes, 26 ; vessels, 94 vascular bundles, 124
;

Orchidaceae, 73, 548 aerial roots, 48

; ; Papaveraceae, 585
climbing roots, 48 ; epiphytes, 213 ; Papaver is capsulae, 587
mucilage, 71, 79 raphides, 78 ; root
; Papaw, Carica papaya
tubers, 48 ;
siliceous bodies, 78 ; Papilionaceae, 605 tendrils, 44
;

velamen, 109 Papillae, 105

Orchis, *48, *548, *549, *550 Parallel-veined leaves, 34, *522
Organic, acids, 80, 225 ; compounds, syn- Paramecium, symbiosis with algae, 234
thesis of, 5 Para nuts, Bertholletia excelsa
Organography, 10 Parasites, 227 ; reduction of shoot in, 27
Orientation, 267 Parastichies, 41
Origanum, 660 Paratonic movements, 272
Ornithocercus, *343 Parenchyma, 99, 112
Ornithogalum, *539, 541 cell wall, 71 ; ;
Parietal placentation, 504
endosperm, *66 Paris, *542, 543
Oruithophilous plants, 308 Parnassia, 593
Ornithopus, 609 Paronychieae, 571
Orobauchaceae, 666 Parsley, Petroselinum
Orobanche, 229, 666, *670 Parsnip, Pastinaca
Orthospermae, 639 Parthenogenesis, 93, 303, 365
Orthostichies, 41 Partial fruits, 519
Orthotropic, 273 Partite leaves, 33
Oryza, 76, *529 starch, 75
; Passiflora, *504, 630
Oscillaria, *338, 339 Passifloraceae, 630
Osmosis, 179, 195, 204 Pastinaca, 641
Osmotic pressure, 180 Paullinia, 620 ;
anomalous thickening,
Osmunda, *448 *143
Osnuindaceae, 449 Payena, 646
Ostrich fern, Struthiopteris Pea, Pisum
Ourouparia, 672, 673 Pectin compounds, 99
Ovule, *472 Pedate leaves, 34
Ovuliferous scale, 495 Pediastritm, *356
Ovum, 514 Pedicnlnris, 231, 666
Oxalic acid, 80 Peireskia, 630
Oxalidaceae, 610 Pelargonium, *610

3 B
738 BOTANY

PeUia, 4-J9 Phlomis, 660

Peltate leaves, 43 Phlox, ,.-,;
Pdligera, 420 Phoenix, 533
Penicillium, *391 selective power. 195
; Phormium, leaf, *184
Pennatae, 349 Phosphorescence, 245
Pentacyclicae, 645 Phosphorus in plants, 187, 188, 189
Pepper, Piper nigrum Photometry, 278
Perforate leaves, 33 Photosynthesis, 216
Perianth, 40 Phototactic movements, 265
Periaxial wood, 144 Phragmidium, 405, *406, *407
Periblem, 157 Phycocyanin, 62
Pericarp, 519 Phycoerythrin, 62, 374
Perichaetium, 433 Phycomycetes, 378
Periclinal walls, 156 Phyllactinia, 390
Pericyle, 118 Phyllocactus, 631
Peridenn, 145 Phylloclades, 25
-
Peridiueae, 342 Phyllode, 45, *162, *163, 210, 212
Peridinivm, *343 Phylogeny, 2, 9 of the internal structure,
;

Peridium of the Uredineae, 404 ;

of 152
Gasteromycetes, 413 Physalis, *520
Perigynous flowers, *505, 506 Physiology, 6, 171
Periodicity in development. 261 Physodes, 58
Periplasm, 441 Physostigma, 609
Perisperm, 473 Physostigmatis semina, 609
Perisporiaceae, 389 Phytelephas, *94, 225, *473. 533; cell

Perisporieae, 390 wall, 71

Peristome, 436, *437, *438 Phyteuma, 677
Perithecium, *390, *394 Phytolacca, vascular bundles, 124
Periwinkle, Vinca Phytolaccaceae, anomalous thickening,
Permanent tissue, 99 *143
Peronospora, 382, *383 Phytopathology, 167
Peronosporeae, 381 .
Phytophthora, 381, *382
Persea, *582, 583 Phytoteratology, 167
Personatae, 661 Picea, *482, *483, *495, 396 ; succini/era,
Peruvian wax palm, Geroxylon andicola 98
Petal, 37 Picrasma, 614
Petasites, 688 Pilobolus, 384 heliotropism, *276
;

Petiole, 31 ; structure, 122 Pilocarpine, 226

Petroselinum, 641 Pilocarpus, 614
Petunia, *661, 663 Pilostyles, *230
Peziza, *392 P^ t o,^? **
1 ^^ Pilularia, *451
Phaeophyll, 62, 368 Pimento,, 634
Phaeosporeae, 368 Pimenta, 634
Phajus, leucoplasts, *75 Pimpinetta, *637, 640, 643
Phalloideae, 415 Pinaceae, 491
Phallus, *414, 415 Pine, Pinus
Phanerogam, growing point, 157 Pinguicida, 237, 668
Phanerogamia, 471 Pinks, Dianthus
Pharbitis, 283, *284 Pinnate leaves, 33
Phascaceae, 436 Pinnularia, *11
Phascum, *423 Pinus, *481, *483, *494, *497 bordered ;

Phaseoleae, 609 pit, *66 medullary ray, *135, 140 ;

;

Phaseolus, 609 growth, 257

;
starch ; sieve-tubes, *66 wood, *130, *132, ;

grains, *74 *136

Phelloderm, 147 Piper, *553 ;
ethereal oil, 79 ;
vascular
Phellogen, 146 bundles, 124
PhUaddphus, 593 Piperaceae, 552 ; epidermis, 109
Plilfunn, 530 Piperinae, 552
Phloem, 112, *141 parenchyma, 113;
; Pistacia, 619
secondary, 137 Pisum, 609 tendrils, 44, *45
;

Phloeoterma, 117 Pith, 117, 118

 INDEX 739

im, seed, *31H Polytoma, *353

Pits,' 65 ; branched, 66 membrane, 65
; Polytomy, 18 false, 18 ;

Pitted vrvs'-ls, !Ci Polytrirliarear, conducting strands, 111

Pix, burgundica, 498 ;
li(|iiida, 498 PatytritJuan, 432, *434, 438
Placenta, 472, 503 Pomeae, 595
Placeutation. *504 Poplar, /'H/III/ a*

PUiu ;<,<-)i ;/, *429, *14 Poppy, J'li/iin-i',-

Plagiotropie, 273 /'"/inltt.'i, *556 ; adventitious shoots, 20 ;

Plankton, 351 bud, *39

I'/n/tktoniella, *350 Poricidal, dehiscence, 519
r
Plantaginaceae; 668 Porogamy, *51. )

Plantagb *511, 668, *670 bundle?, 110 ; ; Positive, geotropism, 280 ; lieliotropism,
protogyny, *310 triplocaulescent, ; 275
28 I'otamogeton, *523
Plantain, Pluittnu" Potamogetouaceae, 524
Plants, distinct from animals, 3 Potassium in plants, 187, 188
Plamodiophora, 342 Potato, Solatium tubi'mxi/m
Plasmodiiim, 54, *55, 341 Potato disease, 381
Plasmolysis, 71, 180 Potentilla, *506, *594, 597
l'lnsiiii'?'i, 382 Potentilleae, 597
Platanauceae, 593 Prickles, 107
Platanus, 593 Primary members, the metamorphosis of,
Platycerium, 251 ; heterophylly, 35 15
I'lec'tridium, *334 Primary meristem, 99 tissue, 100 tissues, ; ;

Pleiochasiuin, 51<> distribution of, 117

Pleomorphism, 163 Primitive form of membrane, 10
Plerome, 157 Primordial, leaf, 31 utricle, 54 ;

Pleurocladia, *368 1'i'i Hilda, glandular hairs, 108 hetero- ;

PltnronffMO, 350 styly, *311 siitenxix, *107 rinensis,

; :

Plumbagiuaceae, 649 <i* \ glandular hairs, *107

Poa, 530 Primulaceae, 647
Podalyrieae, 606 Primulinae, 647
Podophylli rhizoma, 580 Privet, Li<jnxt rum
Podophyllum, 580, *581 Procambium strands, 113, 128
Pm&upora, *394, 395 Products of assimilation, transfer of, 222 ;

I'odostemaceae, 591 adventitious shoots,

; utilisation of, 221
29 emergences, 107
; hapterae. f>l ; ;
Proembryo, 483, 516
roots, 48 Promeristem, 99
Polar, caps, 81 nuclei, 514
; Promyceliuni, 403
Polarity, 250 Prophases, 84
Polemoniaceae, 657 Proaenehyma, 99 -

I'lilemonium, 657 Proserpinaca, growth, 260

Pollen, chamber, 486 grains, 475, *503, ; Protaudry, 310
*513 sacs, 475
; tube, 476, *481, ; Prothallium, *439, 443, *449, 453, *4">4.
*513 tube, rate of growth of, 255
; *457, 458, 460, *463, *405, 473
Pollination, 306 Protococcales, 335
Pollinium, 654 Protogyuy, *310
Polyangium, *337 Protonema, *423, 431
Polyblepharis, *353 Protophloem, 114
Polycarpicae, 571 Protoplasm, 53, 54, 173 ;
circulation of,
Polyembryony, 302 58 ;
inclusions of, 73. ; movements of,
Polygala, *6U, 615 58 ; reactions of, 57 rotation of, 58 ;

Polygalaceae, 614 Protoplast, 53 ;

connection of, 93 ;
*94 ;

Polygonaceae, 567 stipules, 37 ;

differentiation of, 73 ;
movements of,
I'l'iii'innittinii, *f>22, 543 rhizome, *23, ;
265
24 Protoxylem, 114
/'in' I/Hint
inn, *514, *515, *")67 Pmneae, 598
/'(i/if/tui/iitiii, 446. *449, *450; heterophyllv. Pruni virgin ianae cortex, 599
36 I'm a us, *594, *598 ai-ium, bud scales, ;

Polyporeae, 410 *36 cerasus, 511

; cerasus, gum, 71 ;

/'"li/iii'i-i/x, *411 98 ; spinosa, thorns, 28

740 BOTAXY

412 hypha, *60

/suffiota, *411, ; *112; hcterophylly, *:i4, :I5 :
l,-:.f,

335
/'.iflll/IIIIUHHIS, 35
Pseudoparenchyma, 386 ,i, *210, 686
-

/'.ii-i/i/nfftiii/ii, -19;") llaphc, 17 _'

/W///;/,, 633 Itaphides, *77, 7-s

Pt.-ridophyta, 113, 438; bundles, 113, Receptacle, 448

126 fossil, 466
; ; root, 158 second- ;
l!i-il
Algae, Rhodophyoeae
ary growth, 127 Reduced structures, 10
Pteridospermeae, 500 Reduction, division, 84, *85
1't.Tis, *439, *440, 449 ;
concentric Regeneration, 151
bundles, *115 petiole, ;
123 root ; Rejuvenescence, 65
apex, *158 ; vessels. !'ii
Reproduction, 296
Pterocafpi lignum, 609 /, 591 ;
multicellular formation,
Pterocarpus, 609 woo 1. 133 ;

I'lii-i-inia, *404, *405 pleomorphism. ;

H'.i Resedaceae, 591
Pulmonariii, 658 Reserve, cellulose. *473 material, in seed, ;

Pulvinus, 33, 392 *473 ; material, storage of, 224

Pumpkin. Cm-n il,itn /,epo Resin, protective covering of, *209
/'iniini, 634, *635 Resiua, 498
Punicaceae, 634 Resins, 79, 226
Purple Loosestrife, Li/fhi-in/i Respiration, 239, *242 heat produced by. ;

Putrefaction, 233 244 intramolecular. 241 movement

; ;

Pycnidium, *394, 396 of gases in, 244

Pycnoconidia, :\\>t\ Respiratory cavity, 104; coefficient, -Ml ;

Pyrenomycetes, 394 roots, *246

Pyrethri radix, 688 Rest-harrow, O/ioi/i*
Pyrola, 646 Resting period, 262
Pyrolaceae, 646 Reticulate vessels, 95
Pyronema, 389, *393 fertilisation, *392 ; Revolving movement, 282
/',//(, *506, *594, *595 peridenn, *147 ; fili<i/n!'iiteiH(i, 350
Pyxidium, *519, 520 Rhachis of leaf. 3:!

Rhamnaceae, 622
Rhamni Purshiani cortex, 623
Quassia, *613, 614 Rliumnus, *622, 623
Quassiae lignum, 614 Rheotaxis, 266
Querciflorae, 556 Rheotropism, 286
Querrus, *560, *561, *562, *688 bark, ; Rheum, 567, *568, *569
*150 bud scales, :$7; cork, *150 ; ; Rhinantheae, 665
galls, 166 lateiit buds, 23 ;
medul- ; Rhinanthus, 231
lary rays, 141 midsummer growth, ; Rhipidium, *512
132 ; suber, bark, 148 ; saber, cork, is 631
148 ; vessels, 96 tdicicola, 232
Quillaiae cortex, 599 Rhizoids, 15, 50 ;
of Musci. !:;_'

Vin'llaja, 594, *595 Rhizomes, 24

V" i inn-la tendrils, 289 Rhizomorpha, 246
Quince, Cydonin Rhizii/ilioru, 632, *H33
Quinine, 226, 673 Rhizophoraceae, 632
Rhizophores, 50
R!ii-j'i>i<x,
*:!s3 streaming : protojtlasm
Raceme, 509, *510, *511 268
Ilacemose inflorescence, 509, *510 Rhodiiili'itilrn/i. *504, 646
Radial symmetry, 15 of flowers, 507 ; Rhmlophyceae, 373
Radial walls, 156 Rhodoreae, 646
Radium, influence on growth, 259 Rhoeadinae, 585
Radix 567
rhei, Rhoeados petala. 5^7
645 ; reduction of shoot, 27 I'll UN, 619
Rafflesia,
l!atile>iarcae, 645 haustoria, 49 para-
; ;
A' //'.-, *51'2, ;".'.':;

sitism, 229 Riccia, *14, *425

liii, 666 Ricciaceae, 425
Uanunculaceae, 573 Rice, <>/-i/--
Hit a ii<-iil im, *506, *574, *575
it bundle. ; Riclm nl in, 536
 INDEX 741

/'/</// ..-, 017, *61S, *619 ;

aleurone grains, Saffron, 544
*76 palisade cells, 122
; Sage, Salvia
Rigor, cold, heat, etc., 296
Kinged bark, 148 Salicaceae, 555
ll<iliinia, 608 ; adventitious shoots, 20 ; Saliciflorae, 555
leaf, 34 ; movements, 293 :
thorns, Salicin, 556
*45 ; tyloses, *133 Still,;,i-nia, *192 organic acids, 80 ;

Rocella, 418 Salix, *555, 556 adventitious roots, 47;

;

Rontgen Rays, influence on growth, 259 buds, 23 ; polar is, *213

Root, 15, 45, *114, *119, *120, 158;
adventitious, 47 aerial, 46 ; apex of, ; Salsola, organic acids, 80
*158,*159 branching, 46 cap,15,46, ; ;
tut >;<(, *659, 660 pollination, *312 ;

158 central cylinder, 120 contrac-

; ; .^dfiitia, 49, *452, *453, *454, 466
tion of, 323 growth of, 256 -hairs, ; ; Salviniaceae, 452
46, 105, 120, *198, 199 hypoderma, ; Sambuceae, 674
121 lateral, ;
161 lenticels, 150 ; ;
Sambuci flores, 675
mechanical tissues of, *184 meta- ; Sombucus, 674, *676 ; leuticel, *149 ;

morphosis of, 48 parasitic, 231 ; ; stem, *183

phellogen, 149; -pocket, 46; -pressure, Samolus, axillary shoot and leaf, *22
*200, 204 primary cortex, 120 ; ; Sangnisorba, *594, 597
secondary thickening of, 142 -stocks, :
Sanicula, 640
24 structure of, 120 subterranean,
; ; Santalaceae, 644
47 -system, 198
; -system length of, ; Santalin, 133
198; tubers, *48 tubercles, *231, ; Santalum, 644
232 vascular strands of, 127
; Santoninum, 688
Rosa, *520, *594, 595 adventitious ; Sap cavities, 53
shoots, 20; prickles, 107; stipules, Sapindaceae, 620
36 Sapindinae, 618
Rosaceae, 593 Sapona/ria, *571
Rosae Gallicae Petala, 593 Saponin, 80
Rose, Rosa Sapotaceae, 646 gutta-percha, 79 ;

Rose of Jericho, Odontoapenwwm Saprolegnia, *380 ; parthenogenesis, 303

Roseae, 595 Saprolegniaceae, 380
Rosiflorae, 593 Saprophytes, 227, 231
Rosmarinns, 660 Sap wood, 133
Rotation of protoplasm, 58. 267 Sarcina, 336
413
A'"-.//,-.*, Sarcocaulon, resinous covering, *209
Jlubia,673 Sargassum, 367
Rubiaceae, 671 Sarothamnus, 213, 607
Rubidium in plants, 187 Sarracenia, 238, 591
Rubiinae, 668 Sarraceniaceae, 591
lln/ts. *521, *597; adventitious shoots, Sarsaparilla, 543
20 Sassafras, *583 radix, 583 ;

Rudimentary structures, 10 Satureja, 660

llniuf.'-, 567 adventitious shoots, 20
; ; Saxifraga, 593 lime scales, 211
; ;
water
organic acids, 80 stomata, 104
Runners, 25 Saxifragaceae, 592
I!i'l>/>iri, '524 Saxifrage, Saxifraga
Ruscus, cladode, 25, *26 Saxifraginae, 591
1'iifxiil,/. hymenium, *408 s,;ii>;, >.<,!
t
680
Rust fungi, Uredineae Scalariform vessels, 95. *96
linst of wheat, Pun-iniii </////////> Scale hairs, 107 ; -leaves, 31, 37
Ili'fi', *612 Scaly bark, 148
Kutaceae, 611 Scammoniae radix. 657
Rye, fiecale Scandix, 641
356
Sn<vh<i mm i/res, *11, 91, 398, *399 ;
ccll- *435 phosphorescence, *246
, ; ;

liudding, 91 fermentation,
; 243 ; protonema, *176
zymase, 223 Sclii/aeaceae, 449
Saccharomycetes, 398 Schizocarp, *520
tiaccharum, 529, 530 ; wax, *101 Schizogenic intercellular spaces, 98
742 BOTANY

Schizomycetes, 332 Sexual organs of Musci, 432

nemo, 348 Sexual reproduction, 298, 304
Sdiizosaccharv/iiyi-i*. :'.<",<
Shade-loving plants, 176
SchoenocaiUon ,
543 Shoot, 18-29; apex, 18, "19; branching
Schulze's macerating mixture, !'!
of, 19 development, 18 endogenous
; ;

x-;n<t, 541 20; exogenous, 20; origin, 160;

S *525
/-/'*, short, 21 subterranean, 23
; sym :

Scitamineae, 546 metry of, Iti

Sderanthus, 571 Shnmi., 628

Sclerekles, 67, 73, 116 Shrubs, 29
Sclerenchyiiia. 72. 1H>. 182 Sicyos tendril, "289
Sderodeniw, 413, *414 Sieve plates, 67, 93 ; tubes, "66, 93, *95.
Sclerospora, 382 112, 137, 222
Sclerotia, 56, 386 Sigillarieae, 466
Sclerotic cells, 184 *569, 571
Sedopendrium, 446, *448, 449 Sileneae, 571
Scorzonera, 682 vessels, "96 ; Siler, "673
Scotch Fir, Pin us &//<<.>///.> Silica, 71
Serophularia, 665 Siliceous bodies, 78; earth, 192, 351
Scrophulariaceae, 663 ;
albumen crystals. 77 Silicic acid, 133
Scurvy grass, Cochlea r it >
Silicon in plants, 187, 191
Scutellaria, *508 Siliculosae, 590
Scutellum, *527 Siliqua, 519
Scytovema, 420 Siliquosae, 590
Secale, *528 Sil/iliiiini /tii-iiiiafinn, 277
Secondarj', growth of Monocotyledons, 127; Silver h'r, A/>ii'.*

meristem, 99 ; tissue, 100, 127. 152 Silver in plants, 187

Seditm, 213, *592 Simarubaceae, 614
Seed, 473, *474, *475 474 desic- ; coat, ; Simple leaves, 33
cation, 196 dispersion by animals,
; Xiitapis, 590 albae semiua, 591 nigrae
; ;

319 dispersion by water, 319

;
dis- ; semina, 591
semination of, 318 epidermis, 109 ; ; Sinistrorse stem climbers, *283, 284
germination of, 318-320 leaves, 16*2 ;
;
Siphonalsfc 360 fossil, 465 ;

winged, 318 Siphoneaef/60 nuclei, 60 tissue. 97

; ;

Mdijindla, *461, *462, *463 desiccation, ; Siphonocladiales, 359

195 phosphorescence, 247
;
rhizo- ; 359
ir1<idus,

phores, 50, *51 vascular bundles, 113

;
*588
in in,

Selaginellaceae, 461 639, 641

Selection, theory of, 2 Size of cell, 73
Selective power of cells, 194 Sleep movement, nyctitropic movements
Selenium in plants, 187 Sleep positions of leaves, 291
308
Self- fertilisation, Slime fungi, Myxomycetes
Semi-permeable membrane. 179 Smilax, 543, 688
Sempereivum, 213, 592 Smut fungi, Ustilagineae
Senecio, 688 Snowdrop, ('H/n/it/iHs

Senecioneae, 688 Sodium in plants, 187, 191

Senegal radix, 615 Soil, absorptive power of, 1'J'.'

Senna, 602 Solanaceae, 661

Senna, Alexandrina, 605 tiidica, 605 ; Solanin, 80, 226
Sensitive plant, Mimosa 661, *663 ; nujfinn, 661;
Sepal, 37 tubers, 25, "26
Septicidal dehiscence, 519 420
///(/,

Sequoia, 496 686 ,

Serjania, anomalous thickening of, 143, Konthus, 682

*144 Sophoreae, 606
Sesleria, 212 ,sV/,,/.s *594. 59fi
Si--. -He leaves, 32 Soredia, 417
Seta. 436 Siii-i/finiii. stan-h, 76
Sexual generation of, Aiigiosperms, :".!:; ;
Sorus, 448
of Bryophyta, 422 of Gymnospei nun-. ; Spadiciflorae, 530
480 of Pteridophyta, 439
; Spadix, 509
 INDEX 743

is, 410 glutinous, 76 : reserve, 73 ;

sheath,
Sparganiaceae, 530 117, 119 ; transitory, 224
Spartium, 607, 609 shoots, 27 ; Statice, 649
Spatluxlea, exudation of water from, 210 Statolith theory, 274
x/ ft-; /nl a., 571
1
Stele, 117
Spermaphyta, 478 Stellaria, 571
Spermatium, 376, 388 Stellatae, 673
Spermatozoid, 91, 331, *449, *462, *464, Stem, *117, 118 mechanical tissues of,
;

*487 ; 266
cliemotaxis of, *183 of mossplant, 431 succulent.
; ;

Spermogonia, 388, 404, *420 213 -tendrils, 27

;

Sperms, 478 Slemotdtis, *341, 342

S,,lweriu, 395 Stentor, symbiosis with algae, 234
359 , Sterculia, buttress roots, *185
//.'', 118 Sterculiaceae, 623
*390 Stereome, 182
Sphagnaceae, 434 Stereum, 410
Sphagnum, *432, *433 Slicta, 421
Sphenophyllinae, 467 Stigma, 505 movements of, 296
;

Sphenophyllum, 467 Stigmatic fluid, 211

Spike, 509, *510, *511 Stigmatomyces, *399
Spikelet of Gramineae, *526, *527 Stimulus, 174, 273
Spinach, Spinacia Stinging hairs, 105
Spinacia, 75, 568 ; starch, 75 Stipa, 212, 270 leaf, *211 ;

Spindle, fibres, 81, 88 nuclear, 81, *83 ; Stipules, 31, 36

Spiraea, *594 Stolons, 25
Spiraeeae, 594 Stomata, *102, 205, *206, 244 ; develop-
Spiral vessels, 95 ment 104 mechanism of, 206
of, ;

Spirillum, *11, 333 Stoneworts, Characeae

Spirochaete, *11, 336 Stramonii, folia, 663 semina, 663 ;

Spirogyra, *347; cell division, *89 ;

Stratification of cell wall, 65
movements, 267 Stratiotes, 524
Spirotaenia, *344 Strawberry, Fragnti-in
Splint wood, 133 Streaming of protoplasm, *268, 269
Spo/ir/illa, symbiosis with algie, 234 Streptocarpus, 666
Spongy parenchyma, *122 Streptochaeta, 527
S])ontaiieous generation, 5 Streptococcus, 336
Sporangium, 441, *448, 455 dehiscence, ;
Striations of cell wall, *l>4, 65
448 Strickeria, *394
Spores, 304, 441 desiccation, 196 ; Stroma, 395
Sporocarps, 451, *453 Strontium in plants, 187
SjinrniJiiiiil. *384 Strophanthi semina, 651
Sporogonium, 424, 433, 436 Strophanthus, 651, *653
Sporophylls, 38, 441, 458 Structural deviations, 165
Sporophyte, 164 Struthiopteris, 446
Spurge, Kt'i/Jini-liid Strychnine, 226, 649
Spurious fruit, 521 Strychnos, 649, *652
Squill, 543 Style, 505
Stability of the plant body, 178 Styracaceae, 646
Stachys, 659 Styrax, 646 sty rax praeparatus, 593
;

Stalked leaves, 32 Suberin, 70

Stamen, 37. 475, *502 ; movement of, 296; Subcrised cell walls, 69
rate ofgrowth of, 255 Subsidiary cells, 104, 206
Staminodes, 503 Subterranean roots, 47
Wii/iflia, 656 Succisa, *680
Staphisagriae semina, 578 Suction force of transpiring shoots, 205
Staphyliicoccus, 336 Sugar, 79, 202 ; -cane, Sacchun'm
Starch, 73, 195 ; assimilation, 73 ; grains, Sulphur, 79 bacteria, 243, 335 ; in plants,
:

*74, *93 :
grains, action of di;t-t:i^- 187, 188, 189
on, *224 ;
grains as statoliths, 271 : Sumbiil radix, 643
grains, formation in photosynthesis, Sundew, Drosera
r
218 ; grains, sphti-ritic structure, 7. > ; Supporting fibres, 82
744 BOTANY

Xurirella, 350 Thallophyta, 110, 329 conducting ; tracts.

Suspensor, *463, 464 110 form, 11
:
fossil, 465 ;

Swarm spores, *5f>, *265 ; heliotaetic Thallus, 11

movement of, 277 Tliiinniiiliiiin, 385
Sweetbay, Laurv.s nobili* 7V,/, *628
Xii;'rtia, 651 Theine, 226
Symbiosis, 227, 415 77*.
/,'/;/</,/, *417
Symmetry, relations of, 15 ; floral, 507 Thelephoreae, 410
Sympetalae, 645 624, *625, 626
Symphytum, *513, 658 ,
226
Syinpoilium, 18 Thermotropism, 286
Tl<i- x ;,nn, 231, 644
Synapsis, 84, *85 haiistnria. 1
; I'.

Syin arpous gynaeceuin, 503 Thlini;<i,itli tint,;,,, 249

Xi/iieilrit, 350 Thlaspi, 591
Synergidae, 514 Thornapple, Datum
X'triiiya, *649 Thorns, 28
Thuja, 492 ; germination, *161
Tabasheer, 192 Thus americamim, 498
T<iln'rin'Hiiint<iini, 652 Tliymelaeaceae, 631
Tactile hairs, 108 ;
pits, *65, 67 Thymelaeinae, 631
TaeniophyUwn, 48 Thymol, 660
'/''i/i'fes, 687 Thymus, 660
T'luKn-indus, *601, *602, *603, 605 Tilia, 623, *624 ; hast, *141 ; stem, *138;
Tumiis, 544 wood, *138, *139, *140
Tatiacetinn, 688 Tiliaceae, 623
Tannic acid, 563 Ti//ti,,tlxi<i, 49, 197, 213

Tannin, 71, 73, 78, 93, 133 TUletia, 402, *403

Tapetum, 441 Tilletiaceae, 402
Tiiphrina, *398 Tin in plants, 187
Tap-roots, 47 Tissues, 97 mechanical, 182, *183
; pri- ;

Taraxaci radix, 688 mary, 100

secondary, 100, 127
; ;

Taraxacum, 682, *683, *684 ; apogamy, systems, 99 tension of, 181;

93, 518 tissue

; tensions, 181 Titanium in plants, 187
Taxaceae, 489 Tobacco, 663
Tn.i'iiilivin, 494 Tnlypdlopsit, 36f
Taxus, *482, *490, *491 ; vascular system, Tomato, I.i/i-o/H'i'sii-Hiii
124, *125 Tonoplast, 58
Tea, 628 Tooth wort, Lathraea
Teak, 659 Tozzia, 231, 666
Teazel, J)i/i*ti<-n.-< Tracheae, 35, 95
Tecoma, 666 Tracheides, 72, 96, 112, *132, *135 ;

Tectona, 659 fibrous, 72, 135 ; vascular. 72

Teleutospores, 403 Traction fibres, 82
Tellurium in plants, 187 Tradescantia, cell, *59 circulation of ;

Telophase, 84 protoplasm, 59 direct division, *87 ; ;

Temperature, cardinal points of, 176 in- ; hairs, 107 stomata, *103 ;

fluence on growth, 257 Tragacantha, 609

Tendrils, 27, 44, 288, *289 climbers, 287 :
Tragopogon, 682
Teratology, 167 Transfusion cells, 120
Terebinthinae, 610 Transition cells, llf>
Tegiimentary system, 100 Transitory starch, 224
Terminal leaves, 21 Transpiration, 196, 205-209, 212 current, ;

Terminalia, 632 fruit, 320: *202 current, causes of, 203

;

Ternstromiaceae, 627 Transverse heliotropism, 277

Test'tul i Hit ria, 544 Tr,i,,a, 42
Tetracyclicae, 649 Trees, 29 ; age of, 263 ; ferns. 445, *446 ;

TetragonolcHnUf 607 ferns adventitious roots, IS influence ;

Tetraspores, 375 of wind on, *177 ; weeping varieties,

Teucrium, 659 29, 251

T/ialictri'm, 574 apogamy, 93, 518
; Tremella, *400
Thallium in plants, 187 Tremelliueae, 408
 INDEX 745

liit, 357 U.nim, 418

T,-i<-tn, *342 I'stilagineae, 401
Tr/c/n'Cai'li'/t, 656 ("-/itayv, *401 basidium, *402
;

Trichogyne, 376, *377, :388, 4 lit Utriculcffia, 49, 97, 237, 668; bladder,
Trii'/Ki/iiii/ii'/i, *-141t 43 leaf, 43, *44
;

Trichomes, 105 ; internal, 116 Uvae, 623

Tricoccae, 615 Uvae ursi folia, 646
Trifolieae, 607
Trifiiliiini, 007 movements, 293 stipules,
; ; Vaccineae, 646
36 Vacci nium, *645, 646
Triglodiin, 224 Vacuoles, 53 contractile, 56
;

Trigonella, 607 Valerian, Valeric /m

Trijjlaris, 235 Vn!i riana, 675, *676
Triploeaulesct-nt, 28 Valerianae rhizoma, 675
Triticinn, *527, *528 aleuroue, *77
; Valerianella, 675
Tropaeolaceae. 610 Vullisiieria, 21, 524
Trii/ine<ili'iii, 610 cell, *63 ; exudation, ;
Valvate leaves, 38
*209 ;heliotropism, 277 leaf, 43, ; Vmtilla, *549
*209 water stomata, *105
; Vanillin, 70, 226
Trophoplasm, 59 Variation, external causes of, 165 ;
fluctu-
Tropio curvatures, 270 ating, 3 ;
in hybrids, 316 ;
move-
Truffles, 397 ments, 291
7* /'.</"< hud, *39 Vascular, bundles, 111 system, 100, 109; ;

Tuber, *25 closed, 113; termination of, *116

Tuber, *396, 397 Vascular cryptogams, Pteridopkyta
Tuberaceae, 397 Vascular plants, 154
Tubiflorae, 656 Vascular tracheides, 72, 135
Tubuliflorae, 681 Vaucheria, 361, *362
Tulip tree, Lvriodendron Vegetable ivory palm, Phytelephas
Tulipa, *24, 541 Vegetative, cone, 18 reproduction, 297, ;

Turgidity, 178 300

Turgor, movements due to, 291 Veins, 34
Tussilago, *686, 688 Velamen, 48, 120 :
radicum, 109
'['wining plants, 30, 281 Velum, 412
Tylosis, *133 Venation, 34, 110
Typhaceae, 530 Ventilating system, 98
Venus' Fly-trap, Dionaea
Ulex, 607 Veratrine, 226
riliiiiiiiiiin, 501 Veratrinum, 543
Ulmaceae, 563 Vemtrum, 543
Ulmtu, *31, *515, 563, *564 ; adventi- Vfi-liascum, 33, 664, *667
tious shoots, 20 ;.leaf, *31 \'<;-!>,-,<fi, 657, 659
Ulotlirix, *92, 357, *358' Verbenaceae, 658
Ulotrichales, 357 Vernation, 38
Ulva lactuca, *12 VfrtHiica, 213, 665 ; ciqyressoitlex, *212
Umbel, 509, *510, *511 Vci-i-ti.caria, 418
Umbelliferae, 636 oil ducts, 98 ; Vessels, 95 ; pitted, 95 ; reticulate, 95 ;

Umbelliflorae, 635 scalariform, 95, *96 :

spiral, 95
I'lii'tirin. ('<!''<
Vibrio, *333, 336
UncinulH, 390, *391 Vilinrnum, 674
Uniaxial, 28 Vicia, *604, 609 geotropism, *281 ; ;

Unicellular hairs, 107 growth, 257 root tubercles, *231

;

Upper leaf, 31 Vicieae, 609

Uragoga, 672, 673, *674 Victoria regia, heat produced by respira-
Uredineae, 403 tion, 244
Uredospores, 407 \'iitcti, 651, *655 ; sclerenchymatous liinc
(7ri/ini'a, *540, 541 *64
Urticii, 566 hybridisation, *315
; ; stinging Viiicftaxicum, 654, *655
hairs, 105. *106 Vine, Vifi.t

Urticaceae, 566 V!!,', *188, *507, *519, *629, 6:50 ;

Urticinae, 563 rmiTgences, 107, *108; papilla-, *\Q[>

740 110TANY

Violaceae, 629 Wljli<i, 49, 197, 53

Violet, Vinlii Wood, autiiinn, 131 ;
elements, *137 :

Visi-in-in, *f,ii'i fibres, 134, 136; parenchyma. 1-">I.

\'in<- ntn, *94, 231, *644 ; i-ptdt-rmis, 1-1(5 ; 140; secondary. l :

''t :
spring, 131
false iliubotomy, 18 Woodrull', Ax/iei-iiln
Vitai-L-ae, 623 ; iiluiiiiniuiii, 192 Wormwood, .1 //<//(/.</(/

Vital force, 174 Wounds, 1i,-alin- of, 150

Vital properties of plants, 172
\'itis, *622, 623 ; tendrils. 27
Xant lieiii, 80
Vittae, 98
Xanthophyll, 62, 63
Fo/teifl, 501
Xanthoprotein, reaction, 57
Volva, 412
Xmithoria, *416, *417
Volvocales, 353
Xenia, 305
Volrox, *354, 355
Xerophytes, 212; stomata of, 104
Xylem, 111, 112; jiari-ncliyina. 112
\Vnli-liia, 500
Xylocliromc. ]'>">
Walking fern, Camptosorus rhizophyllna
Water, absorption of, 197 constitution, ;

196 -culture, *190

;
distribution of, ;
Yam, Itio
200 ; exudation of, 209 ; imbibition, Yew, Taxus baccata
194 Yin-,-,,, *511, 543; pollination, 308;
necessary for
; assimilation,
219 245 -plants, growth in thickening, 144
; -plants, ;

air and water, 260; -stomata, 104 ;

*105, 123 -supply, regulation of,

; Zinnia, 485, *487, *488 :
spermatozoid,
212 *487
Water Crowfoot, fianuncttlus at/ti/i.-< Zaitichellia, 524
Water-ferns, Hydropterideae Zea, 528; bundles, *110, *111 exuda- ;

Water Hemlock, Cicuta tion of water from, 209 stem, *117 ;

Water-net, Hydrodiciyon Zinc in plants, 187

Water Parsnip, Stum Zin (jiber, 546, *547 ethi-n-al oil, 79
;

Water-pores, water-stomata Zingiberaceae, 456

Wax, *101 Zinnia, 687
Weigelia, 675 Zostera, 524 ; pollination, 307
\\'f/i'-itschia, 498
:>> J'>
Zygnemaccae,
Wheat, Triticum Zygomorphic, flowers, *508 ; symmetry,
White Thorn, 16
WiHovghbeia, 651 Zygomycetes, 383 >^__
Willow, Salix Zygophyllaceae, 611
Willow-herb, ZygoMceharotnyces, 399
Winter buds, 109 Zygospore, 331, 344, 3S4, *3S5
Witches'-brooms, 398 Zymase, 223

THE END

rrintedby K. & R. CI.AKK, I.lMiTFti. Edinburgh.

 8vo. 18s. net.

THE ORIGIN OF A LAND FLORA

A THEORY BASED UPON THE FACTS OF
ALTERNATION

BY

F. 0. BOWER, Sc.D., F.R.S.

REUU'S PROFESSOR OF BOTANY IN THE UNIVERSITY OF GLASGOW

With numerous Illustrations.

In his Preface the author explains that his interest in the question dis-
cussed in the presentwork has been of long standing that it seemed to him
;

probable that some biological cause had determined the prevalence and con-
stancy of the alternation, to which apogamy and apospory appear as occasional
exceptions. The theory was entertained that the change of conditions
involved in the invasion of the land by organisms originally aquatic had

played a prominent part in the establishment of those alternating phases of

the life-cycle which are so characteristic of Archegoniate plants. As early as
1890 Professor Bower briefly stated his theory in a paper published in the
Annals of Botany, but his researches have led him to the conclusion that only
"
now, after the lapse of seventeen years, it has been possible to state the
biological argument more fully in the present volume, strengthened by many
new facts. The first part deals with the general theory. The second part is

taken up with a detailed statement of the facts, together with comparison of

the constituents of the several phyla inter se. The third part is devoted to
general comparisons and conclusions. . . .

" The
attempt has been made to work in the results of Palaeontological
research with those of the comparative analysis of living forms. The enquiry
has related to all the characters, both vegetative and propagative, of the

sporophyte generation these include the external form, the embryogeny,

:

and anatomical features, and especially the structure and development of the
spore-producing members, while the characters of the gametophyte have also
been taken into account. It is found that the conclusions arrived at are

supported by general convergence of the lines of evidence derived from all of

these sources."
Professor Bower's important work is a substantial volume of over seven
hundred pages, with a large number of illustrations.

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