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 1. Anatomy is the study of the body’s structures.
• Developmental anatomy considers anatomical changes from conception to adulthood.
Embryology focuses on the first 8 weeks of development.
• Cytology examines cells, and histology examines tissues.
• Gross anatomy studies organs from either a systemic or a regional perspective.
2. Surface anatomy uses superficial structures to locate deeper structures, and anatomical
imaging is a noninvasive technique for identifying deep structures.
3. Physiology is the study of the body’s functions. It can be approached from a cellular or a
systems point of view.
4. Pathology deals with all aspects of disease. Exercise physiology examines changes
caused by exercise.
1.2 Structural and Functional Organization of the Human Body
1. Basic chemical characteristics are responsible for the structure and functions of life.
2. Cells are the basic structural and functional units of all living organisms. Organelles are
small structures within cells that perform specific functions.
3. Tissues are composed of groups of cells of similar structure and function and the
materials surrounding them. The four primary tissue types are epithelial, connective,
muscle, and nervous tissues.
4. Organs are structures composed of two or more tissues that perform specific functions.
5. Organs are arranged into the 11 organ systems of the human body (see figure 1.3).
6. Organ systems interact to form a whole, functioning organism.
1.3 Characteristics of Life
Humans share many characteristics with other organisms, such as organization,
metabolism, responsiveness, growth, development, and reproduction.
1.4 Biomedical Research
Much of our knowledge about humans is derived from research on other organisms.
1.5 Homeostasis
Homeostasis is the condition in which body functions, body fluids, and other factors of the
internal environment are maintained at levels suitable to support life.
Negative Feedback
1. Negative-feedback mechanisms maintain homeostasis.
2. Many negative-feedback mechanisms consist of a receptor, a control center, and
an effector.
Positive Feedback
1. Positive-feedback mechanisms usually increase deviations from normal.
2. Although a few positive-feedback mechanisms normally exist in the body, some
positive-feedback mechanisms are harmful.
3. Normal positive-feedback mechanisms include blood clotting and childbirth
labor. Harmful positive-feedback examples include decreased blood flow to the
heart.
1.6 Terminology and the Body Plan
Body Positions
1. A human standing erect with the face directed forward, the arms hanging to the sides,
and the palms facing forward is in the anatomical position.
2. A person lying face upward is supine; a person lying face downward is prone.
Directional Terms

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written consent of McGraw-Hill Education.
2
 Directional terms always refer to the anatomical position, no matter what the actual
position of the body (see table 1.2).
Body Parts and Regions
1. The body can be divided into a central region, consisting of the head, neck, and trunk,
and the upper limbs and lower limbs.
2. Superficially, the abdomen can be divided into quadrants or into nine regions. These
divisions are useful for locating internal organs or describing the location of a pain or
a tumor.

Planes
1. Planes of the Body
• A sagittal plane divides the body into right and left parts. A median plane divides
the body into equal right and left halves.
• A transverse (horizontal) plane divides the body into superior and inferior
portions.
• A frontal (coronal) plane divides the body into anterior and posterior parts.
2. Sections of an Organ
• A longitudinal section of an organ divides it along the length of the organ.
• A transverse (cross) section cuts at a right angle to the length of the organ.
• An oblique section cuts across the length of the organ at an angle other than a
right angle.
Body Cavities
The body contains two types of internal cavities: the dorsal and ventral body cavities.
1. The dorsal body cavity contains the cranial cavity and the vertebral canal.
2. The ventral body cavity houses the thoracic cavity and the abdominopelvic cavity.
The mediastinum subdivides the thoracic cavity.
3. The diaphragm separates the thoracic and abdominal cavities.
4. Pelvic bones surround the pelvic cavity.
Serous Membranes
1. Serous membranes line the trunk cavities. The parietal portion of a serous membrane
lines the wall of the cavity, and the visceral portion is in contact with the internal
organs.
• The serous membranes secrete fluid, which fills the space between the visceral
and parietal membranes. The serous membranes protect organs from friction.
• The pericardial cavity surrounds the heart, the pleural cavities surround the lungs,
and the peritoneal cavity surrounds certain abdominal and pelvic organs.
2. Mesenteries are parts of the visceral peritoneum that hold the abdominal organs in
place and provide a passageway for blood vessels and nerves to the organs.
3. Retroperitoneal organs are located “behind” the parietal peritoneum.

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written consent of McGraw-Hill Education.
3
Topics Related to the Study of Anatomy and Physiology
The use of animals in research is relevant, and the students may have strong opinions about the
ethical issues involved. Discuss pros and cons (including financial considerations) for
alternatives to animal experimentation, such as tissue culture and computer simulation.

Anatomical anomalies can be used for discussion concerning the concept of normal. Anatomy
and physiology are replete with references to normal and abnormal structures and values.
Students will benefit from the clarification of the meaning of the word “normal" as it will be
used within the context of the course.

Newspaper, magazine, or internet sources related to the new imaging technologies can help
students appreciate the amount of knowledge of anatomy and physiology a diagnostician must
possess in order to interpret those potentially meaningless images. Use table 1.1, Anatomical
Imaging, as the starting point for a homework assignment to find out more information.

There are excellent photographs throughout the text that illustrate technological advances in
imaging techniques. The advents of the electron microscope, patch-clamping, micro-electrodes,
and radio-immunoassay have increased our ability to investigate cell structures and cell
membrane transport. The newest scanning tunneling electron microscopes have taken resolution
down to the level of individual molecules. Class discussion could focus on the intriguing area of
cellular research.

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written consent of McGraw-Hill Education.
4
 Themes in Chapter 1
Structure and Function

Medical Terminology
“When in Rome…” is a concept that could be applied to knowing and using anatomical and
medical terminology. Students must use their language in order to communicate with other
scientists and healthcare professionals. Students need to learn that there is value in the
precision of anatomical terminology. The notion that the body is a collection of interlocking
parts is a concept foreign to many students, who view the body as a singular and solid entity.
Students may not realize there is a connection between the words that are used in class and
their own bodies. Point out the valuable list of prefixes, suffixes, and combining forms on the
back cover of the book and the Glossary (pages G-1 to G-3) that will help them gain a mastery
of this “new” language. Also useful is Table 1.2, Directional Terms for Humans.

Homeostasis

Feedback
Spend time on the concepts of positive and negative feedback to ensure student understanding.
Provide examples in addition to those provided in the text. Ask students to think about and
then discuss examples of events that push the body out of homeostasis and how the body
returns to homeostasis. Discuss ways the body can be helped to return to homeostasis in
emergencies. Be sure students understand how to interpret the Process Figure 1.4, Negative-
Feedback Mechanism: Body Temperature, and Homeostasis Figure 1.5, Negative-Feedback
Control of Body Temperature, because this format is used throughout the book and can be an
invaluable tool in understanding complex body processes.

Cell Theory and Biochemistry

Students must assimilate this foundational knowledge before they can grasp more complex
physiological processes like cell membrane transport and cell-to-cell communication. Stress
the pivotal position of cells and biochemistry in understanding higher levels of organization.

Changes Through Time

Students must grasp the difference between structures/parts and functions/processes.

Introduce the element of time and the possibility of change through time (moment to moment,
over the life span, and evolutionarily) in both structures and functions.

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written consent of McGraw-Hill Education.
5
 Learning Outcomes Correlation with Predict Question Types
Question Type Question # Bloom's level Learning Outcome
Learn to Predict Application 1.5a,b
Predict 1 Evaluation 1.5a,b
Predict 2 Evaluation 1.2a
Predict 3 Evaluation 1.5a,b
Predict 4 Comprehension 1.5b
Predict 5 Comprehension 1.5b,c
Predict 6 Comprehension 1.6a,b
Predict 7 Comprehension 1.6b
Predict 8 Comprehension 1.6f,g,h

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written consent of McGraw-Hill Education.
6
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to this creature of thought, and to conceive of the type in the sense
of a Platonic idea. This was the only view that was possible in
combination with the dogma of the constancy of species, and so
Elias Fries, in his ‘Corpus Florarum,’ 1835, in speaking of the natural
system, could consistently say, ‘est quoddam supranaturale,’ and
maintain that each division of it ‘ideam quandam exponit.’ So long as
the constancy of species is maintained, there is no escaping from
the conclusion drawn by Fries, but it is equally certain that
systematic botany at the same time ceases to be a scientific account
of nature. Systematists, adopting this conclusion as necessarily
following from the dogma, might consider themselves as seeking to
express in the natural system the plan of creation, the thought of
the Creator himself; but in this way systematic botany became
mixed up with theological notions, and it is easy to understand why
the first feeble attempts at a theory of descent encountered such
obstinate, nay, fanatical opposition from professed systematists, who
looked upon the system as something above nature, a component
part of their religion. And if we look back we find that these views
are based on the dogma of the constancy of species, while Linnaeus’
‘Philosophia Botanica’ teaches us on what grounds this dogma rests,
where it says, ‘Novas species dari in vegetabilibus negat generatio
continuata, propagatio, observationes quotidianae, cotyledones.’
In spite of all this one important advance was made by the
successors of Jussieu; the larger groups of genera, the families,
were defined with the certainty and precision, with which Linnaeus
had fixed the boundaries of species and genera, and were supplied
with characteristic marks. They succeeded also in clearly
distinguishing various still larger groups founded on natural affinity,
such as the Monocotyledons and Dicotyledons; the distinction
between Cryptogams and Phanerogams was by degrees better
appreciated, though this point could not be finally settled, so long as
it was attempted to reduce the Cryptogams entirely to the scheme of
the Phanerogams. The chief hindrance however to the advance of
systematic botany, at least at the beginning of this period, lay in the
defective morphology enshrined in Linnaeus’ terminology and in his
doctrine of metamorphosis. A great improvement certainly was
effected in the early part of the 19th century by De Candolle’s
doctrine of the symmetry of plants,—a doctrine which has been
much undervalued, and that merely on account of its name; it is
really a comparative morphology, and the first serious attempt of the
kind since the time of Jung that has produced any great results; a
series of the most important morphological truths, with which every
botanist is now conversant, were taught for the first time in De
Candolle’s doctrine of symmetry in 1813. But one thing was wanting
not only in Jussieu and De Candolle, but in all the systematists of
this period, with the single exception of Robert Brown, and this was
the history of development. The history of the morphology and
systematic botany of this period shows indeed, that the comparison
of mature forms leads to the recognition of many and highly
important morphological facts; but as long as matured organisms
only are compared, the morphological consideration of them is
always disturbed by the circumstance that the organs to be
compared are already adapted to definite physiological functions,
and thus their true morphological character is often entirely
obscured; on the other hand, the younger the organs are, the less is
this difficulty experienced, and this is the real reason why the history
of development is of so great service to morphology. It was then one
of the characteristic features of the period we are describing, that its
morphology was formed upon the study of matured forms; the
history of development, or at all events of very early stages of
development, could not be turned to account till after 1840, for skill
in the use of the microscope, here indispensable, was not sufficiently
advanced before that time to make it possible to follow the growth
of organs from their first beginnings.
The establishment of natural affinities combined with the
assumption of the constancy of species, the growth of comparative
morphology without the history of development, lastly, the very
subordinate attention still paid to the Cryptogams,—these are the
special characteristics of the period which has now to be described
at greater length.
 Here we must once more call attention to the fact, that Linnaeus
was the first to perceive that a system which was to be the
expression of natural affinities could not be attained in the way
pursued by Cesalpino and his immediate successors. All who have
attentively studied the writings of Linnaeus which appeared after the
‘Classes Plantarum’ (1738) must have seen the difference between
that way and the one recommended by him—a difference which is
the more obvious because Linnaeus himself, like his predecessors,
constructed an artificial system on predetermined principles of
classification, and always employed it for practical purposes, while
he published at the same time in the above-named work his
fragment of a natural system, and in the preface set forth the
peculiar features of the natural and artificial systems in striking
contrast with one another. The first thing and the last, he says in his
prefatory remarks to his fragment, which is demanded in systematic
botany, is the natural method, which slighted by less learned
botanists has always been highly regarded by the more sagacious,
and has not yet been discovered. If, he continues, we collect the
natural orders from all existing systems (up to 1738), we shall get
but a small list of really allied plants, though so many systems have
claimed to be natural. He had himself long laboured to discover the
natural method and had found out some things that were new; but
though he had not succeeded in carrying it through to a perfect
work, he would continue his efforts as long as his life lasted. He
makes the very important remark, that a key, that is, a priori
principles of classification, cannot be given for the natural method,
till all plants have been reduced to orders; that for this no a priori
rule is of value, neither this nor that part of the fructification, but the
simple symmetry alone (simplex symmetria) of all the parts, which is
often indicated by special marks. He suggests to those who are bent
on trying to find a key to the natural system, that nothing has more
general value than relative position, especially in the seed, and in
the seed especially the ‘punctum vegetans,’—a distinct reference to
Cesalpino. He says that he establishes no classes himself, but only
orders; if these are once obtained, it will be easy to discover the
classes. The essence of the natural system could not have been
more clearly expounded in Linnaeus’ time, than it is in these
sentences. He established as early as 1738 sixty-five natural orders,
which he at first simply numbered; but in the first edition of the
‘Philosophia Botanica’ in 1751, where the list is increased to sixty-
seven, he gave a special name to each group; and he showed his
judgment by either taking his names from really characteristic
marks, or what was still better, by selecting a genus and so
modifying its name as to make it serve as a general term for a whole
group. Many of these designations are still in use, though the extent
and content of the groups have been greatly changed. This mode of
naming is an important point, because it expresses the idea, that the
different genera of such a group are to some extent regarded as
forms derived from the one selected to supply the name. Many of
Linnaeus’ orders do in fact indicate cycles of natural affinity, though
single genera are not unfrequently found to occupy a false position;
at all events, Linnaeus’ fragment is much the most natural system
proposed up to 1738, or even to 1751. It is distinguished from
Kaspar Bauhin’s enumeration in this, that its groups do not run into
one another, but are defined by strict boundaries and fixed by
names.
The Linnaean list is distinctly marked by the endeavour to make
first the Monocotyledons, then the Dicotyledons, and finally the
Cryptogams follow one another; that the old division into trees and
herbs already rejected by Jung and Bachmann, but still maintained
by Tournefort and Ray, disappears in Linnaeus’ natural system will be
taken for granted after what has been already said of it, and from
this time forward this ancient mistake is banished for ever.

In Bernard de Jussieu’s[34] arrangement of 1759 we find some

improvements in the naming, the grouping, and the succession, but
at the same time some striking offences against natural affinity. He
published no theoretical remarks on the system, but gave expression
to his views on relations of affinity in the vegetable kingdom in
laying out the plants in the royal garden of Trianon, and in the
garden-catalogue. His nephew published his uncle’s enumeration in
the year 1789 in his ‘Genera Plantarum,’ affixing the date of 1759
given above. The difference between it and the Linnaean fragment
does not seem sufficiently marked to make it necessary to reproduce
it here. It should be noticed however that Jussieu begins with the
Cryptogams, passes through the Monocotyledons to the
Dicotyledons, and ends with the Conifers. Adanson’s claims of
priority over Bernard de Jussieu (see the ‘Histoire de la Botanique’
de Michel Adanson, Paris, 1864, p. 36) may be passed over as
unimportant. The natural system was not advanced by Adanson to
any noticeable extent; how little he saw into its real nature and into
the true method of research in this department of botany is
sufficiently shown by the fact, that he framed no less than sixty-five
different artificial systems founded on single marks, supposing that
natural affinities would come out of themselves as an ultimate
product,—an effort all the more superfluous, because a
consideration of the systems proposed since Cesalpino’s time would
have been enough to show the uselessness of such a proceeding.
The first great advance in the natural system is due to Antoine
Laurent de Jussieu[35] (1748-1836). After all that has been said no
further proof is needed that he was no more the discoverer or
founder of the natural system than his uncle before him. His real
merit consists in this, that he was the first who assigned characters
to the smaller groups, which we should now call families, but which
he called orders. It is not uninteresting to note here how Bauhin first
provided the species with characters, and named the genera but did
not characterise them, how Tournefort next defined the limits of the
genera, how Linnaeus grouped the genera together, and simply
named these groups without assigning to them characteristic marks,
and how finally Antoine Laurent de Jussieu supplied characters to
the families which were now fairly recognised. Thus botanists learnt
by degrees to abstract the common marks from like forms; the
groups thus constituted were being constantly enlarged, and an
inductive process was thus completed which proceeded from the
individual to the more general.
It might appear that the merit of Antoine de Jussieu is rated too
low, when we praise him chiefly and simply for providing the families
with characters; but this praise will not seem small to those who
know the difficulty of such a task; very careful and long-continued
researches were necessary to discover what marks are the common
property of a natural group. Jussieu’s numerous monographs show
with what earnestness he addressed himself to the task; and it must
be added, that he was not content simply to adopt the families
established by Linnaeus and by his uncle and the limits which they
had assigned to them, but that he corrected their boundaries and in
so doing established many new families, and was the first who
attempted to distribute these into larger groups, which he named
classes. But in this he was not successful. His attempt to exhibit the
whole vegetable kingdom in all its main divisions, to unite the
classes themselves into higher groups, was also unsuccessful, for
these larger divisions remained evidently artificial. The three largest
groups on the contrary, into which he first divides the world of
plants, the Acotyledons, Monocotyledons, and Dicotyledons are
natural; but they had been already partly marked out by Ray,
afterwards by Linnaeus, and finally in Bernard de Jussieu’s
enumerations. Still it is the younger Jussieu’s great and abiding
merit, to have first attempted to substitute a real division of the
whole vegetable kingdom into larger and gradually subordinate
groups for mere enumerations of smaller co-ordinated groups,—an
undertaking which Linnaeus expressly declared to be beyond his
powers. If then Jussieu’s system was far from giving a satisfactory
insight into the affinities of the great divisions of the vegetable
kingdom, yet it opened out many important points of view, from
which they could afterwards be discovered, and it certainly became
the foundation for all further advance in the natural method of
classification; for this reason it is necessary to give a view of it in the
following table:—
 A. L. de Jussieu’s System of 1789.
CLASS.
Acotyledones I.
Monocotyledones
Stamina
hypogyna II.
perigyna III.
epigyna IV.
Dicotyledones.
Apetalae Stamina
hypogyna V.
perigyna VI.
epigyna VII.
Monopetalae Corolla
hypogyna VIII.
perigyna IX.
epigyna
antheris
connatis X.
distinctis XI.
Polypetalae Stamina
epigyna XII.
hypogyna XIII.
perigyna XIV.
Diclines irregulares XV.

This table shows that Jussieu did not oppose the Cryptogams,
which he calls Acotyledones, to the whole body of Phanerogams, as
Ray did under the name of Imperfectae; he rather regards the
Acotyledones as a class co-ordinate with the Monocotyledones and
Dicotyledones; but this mistake or similar mistaken views run
through all systematic botany up to 1840; the morphology founded
by Nägeli and by Hofmeister’s embryological investigations first
showed that the Cryptogams separate into several divisions, which
co-ordinate with the Monocotyledons and Dicotyledons. At the same
time the use of the word Acotyledones for Linnaeus’ Cryptogams
shows that Jussieu overrated the systematic value of the cotyledons,
because, as is seen from the introduction to his ‘Genera Plantarum,’
he was quite in the dark on the subject of the great difference
between the spores of Cryptogamic plants and the seeds of
Phanerogams. His conception of the organs of generation was
essentially that of Linnaeus; hence he judged of the Cryptogams
according to the scheme of the Phanerogams, and, not perceiving
their peculiarities, he virtually characterised them by negative marks.
If we notice in the above table how the Phanerogams are
separated into classes, it strikes us that the triple division into
hypogynous, perigynous, and epigynous is repeated no less than
four times; this shows that Jussieu had mistaken ideas of the value
of these marks for classification, whereas the recurrence of them so
often should of itself have suggested a doubt on this point. To judge
of his system more exactly we must here give his series of the
families, which he had already raised to the number of a hundred.

Class I.
1. Fungi.
2. Algae.
3. Hepaticae.
4. Musci.
5. Filices.
6. Naiades.

Class II.
7. Aroideae.
8. Typhae.
9. Cyperoideae.
10. Gramineae.

Class III.
11. Palmae.
12. Asparagi.
13. Junci.
14. Lilia.
15. Bromeliae.
16. Asphodeli.
17. Narcissi.
18. Irides.

Class IV.
19. Musae.
20. Cannae.
21. Orchides.
22. Hydrocharides.

Class V.
23. Aristolochiae.

Class VI.
24. Elaeagni.
25. Thymeleae.
26. Proteae.
27. Lauri.
28. Polygoneae.
29. Atriplices.

Class VII.
30. Amaranthi.
31. Plantagines.
32. Nyctagines.
33. Plumbagines.

Class VIII.
34. Lysimachiae.
35. Pediculares.
36. Acanthi.
37. Jasmineae.
38. Vitices.
39. Labiatae.
40. Scrophulariae.
41. Solaneae.
42. Borragineae.
43. Convolvuli.
44. Polemonia.
45. Bignoniae.
46. Gentianeae.
47. Apocyneae.
48. Sapotae.

Class IX.
49. Guajacanae.
50. Rhododendra.
51. Ericae.
52. Campanulaceae.

Class X.
53. Cichoraceae.
54. Cinarocephalae.
55. Corymbiferae.

Class XI.
56. Dipsaceae.
57. Rubiaceae.
58. Caprifolia.

Class XII.
59. Araliae.
60. Umbelliferae.

Class XIII.
61. Ranunculaceae.
62. Papaveraceae.
63. Cruciferae.
64. Capparides.
65. Sapindi.
66. Acera.
67. Malpighiae.
68. Hyperica.
69. Guttiferae.
70. Aurantia.
71. Meliae.
72. Vites.
73. Gerania.
74. Malvaceae.
75. Magnoliae.
76. Anonae.
77. Menisperma.
78. Berberides.
79. Tiliaceae.
80. Cisti.
81. Rutaceae.
82. Caryophylleae.

Class XIV.
83. Sempervivae.
84. Saxifragae.
85. Cacti.
86. Portulaceae.
87. Ficoideae.
88. Onagrae.
89. Myrti.
90. Melastomae.
91. Salicariae.
92. Rosaceae.
93. Leguminosae.
94. Terebinthaceae.
95. Rhamni.

Class XV.
96. Euphorbiae.
97. Cucurbitaceae.
98. Urticae.
 99. Amentaceae.
100. Coniferae.

Jussieu’s division of the Cryptogams and Monocotyledons offers

much that is satisfactory, if we put the position of the Naiades out of
sight. The grouping of the Dicotyledons on the contrary is to a great
extent unsuccessful, chiefly owing to the too great importance which
he attached to the insertion of the parts of the flowers, that is, to
the hypogynous, perigynous, and epigynous arrangement. It is in
this grouping of families into classes that the weak side of the
system lies; it is utterly artificial, and the task of his successors has
been to arrange the families of the Phanerogams, which were most
of them well-established, and especially those of the Dicotyledons, in
larger natural groups. But this could not be effected, till morphology
opened new points of view for systematic botany; Jussieu, as has
been already remarked, accepted Linnaeus’ views of the morphology
of the organs of fructification in Phanerogams, though he introduced
many improvements in details. He laid greater stress on the number
and relative positions of the different parts of the flower; attention to
their insertion on the flowering axis, which he designated as
hypogynous, perigynous, and epigynous, would have been a great
step in advance, if he had not overrated its systematic value. The
morphology of the fruit is very superficial in Jussieu; even the
designation of dry indehiscent fruits as naked seeds recurs in his
definitions, though as it happens this misconception does not cause
any great disturbance. How inexact was his investigation of the
organs of fructification, when they were somewhat small and
obscure, is best shown by the fact that the Naiades, which are made
to include Hippuris, Chara, and Callitriche, appear among the
Acotyledons, and that Lemna and the Cycads are placed with the
Ferns.
Jussieu explained the dictum, ‘Natura non facit saltus,’ to mean
that the whole body of plants in its natural arrangement must exhibit
a lineal series ascending from the most imperfect to the highest
forms; but he does not say whether Linnaeus’ comparison of the
natural system to a geographical map, the countries in which answer
to orders and classes, is also admissible.
His theoretical observations on the value to be given to certain
marks in a systematic point of view are not attractive, and for the
most part not very correct; he speaks as though some marks must
have a more extensive, others a less extensive value; the perception
of the fact, so far as it is true, rests entirely upon induction; that is,
after the natural affinities have been already recognised to a certain
extent, it becomes apparent that certain marks remain constant in
larger or smaller groups; the systematist can now go on to try
whether such constant marks occur in other plants also, which he
had hitherto assigned to other groups, and thus put it to the test
whether those marks may not be accompanied by others, which
would serve to establish the affinities; that Jussieu did so proceed in
defining his families admits of no doubt, but he was not himself
thoroughly conscious of the fact; at all events, he did not extend this
mode of proceeding, the seeking after leading marks, to the
establishing of larger groups or classes, for these he founded on
predetermined principles.
Jussieu’s labours as a systematist were not confined to the
publication of his ‘Genera Plantarum’; on the contrary, his most
fruitful researches began after 1802, and were continued to the year
1820, and their results appeared in a long series of monographs on
different families in the Mémoires du Museum. He felt with De
Candolle, Robert Brown, and later systematists, that the perfecting
of the natural system depended mainly on the careful establishing
and defining of families. His efforts received a new impulse from the
work of a German writer, whose first volume had appeared in 1788,
a year therefore before the ‘Genera Plantarum,’ a second following it
in 1791, and a supplementary volume in 1805.

This work was Joseph Gärtner’s[36] ‘De fructibus et seminibus

plantarum,’ in which the fruits and seeds of more than a thousand
species are described and carefully figured. But almost more
important than these numerous descriptions, though they offered
rich material to the professed systematists, were the introductions to
the first two volumes, and especially to those of 1788. They contain
valuable reflections on sexuality in plants,—a subject which had
remained in the condition in which it was left by Camerarius (1694)
till it was greatly developed by Koelreuter after 1761, and had since
then been little studied,—and an account of the morphology of fruits
and seeds, the knowledge of which had gone back rather than
advanced since the days of Malpighi and Grew. Gärtner was well
qualified for this work by his unparalleled knowledge of the forms of
fruits, and still more by the character of his mind. Free from
Linnaeus’ scholastic bias, he addressed himself to the examination of
the most difficult organs of plants with as great freedom from
prepossessions as exact acquaintance with the writings of others; he
gives us the impression of a modern man of science more than any
other botanist of the 18th century, with the exception of Koelreuter.
He knew how to communicate with clearness of language and
perspicuity of arrangement whatever he gathered of general
importance from each investigation. Though it is easy to see that the
founding of the natural system was ever before his mind as the final
object of his protracted labours, he was in no eager haste to reach
it; he contented himself with arranging his fruits, saying expressly
that the natural system would never be founded by these means
alone, though the exact knowledge of fruits and seeds supplied the
most important means for decision. Thus his great work was at once
an inexhaustible mine of single well-ascertained facts, and a guide to
the morphology of the organs of fructification and to its application
to systematic botany. The imperfections, which are to be found even
in this work, are due to the circumstances of the time; in spite of
Schmeidel’s and Hedwig’s researches into the Mosses there was still
the old obscurity with regard to the organs of propagation in the
Cryptogams, and this rendered a right definition of the ideas, seed
and fruit, extremely difficult. But Gärtner made one great step in
advance on this very point when he showed that the spores of the
Cryptogams were essentially different from the seeds of
Phanerogams, with which they had been hitherto compared,
because they contain no embryo; he called them therefore not
seeds, but gemmae. The second great hindrance to a true
conception of certain characters in fruits and seeds on the part of
Gärtner was the entire ignorance of the history of development
which then reigned; yet even here we see an advance, if only a
small one, made by him in his repeatedly going back to the young
state for a more correct idea of the organs.
Above all, Gärtner put an end to the blunder of regarding dry
indehiscent fruits as naked seeds, by rightly defining the pericarp as
in all cases the ripened wall of the ovary, and by considering its
strong or weak construction, its dry or pulpy condition, as a
secondary matter. It is obvious that the whole theory of the flower
was thus placed upon a better basis, since dry indehiscent fruits may
come from inferior or superior ovaries. But Gärtner’s theory of the
seed is one of his most valuable contributions to the science. After
careful consideration of the seed-envelopes, he submitted the inner
portion (nucleus) enclosed by them to a searching comparative
examination; he correctly distinguished the endosperm from the
cotyledons, and described the variations in its form and position.
This was the more needful, since Linnaeus had denied the existence
of an ‘albumen’ in plants, which Grew had already recognised and so
named; to Linnaeus it appeared to be of no use to the seed. Though
Gärtner speaks of the cotyledons as uniting with the embryo to form
the nucleus of the seed, yet his account shows that he regarded
them as outgrowths of the embryo itself. The uncertainty which still
existed in the interpretation of the parts of the seed is shown even
in Gärtner by his curious notion of a ‘vitellus,’ which in fact takes in
everything that he was unable to explain aright inside the seed; for
instance, he makes the scutellum in grasses, and even the
cotyledonary bodies of Zamia a vitellus, and applies the same name
to the whole contents of the spores of Seaweeds, Mosses, and
Ferns. In spite of the striking defects connected with this mistaken
notion in his theory of the seed, his views far surpass in clearness
and consistency all that had hitherto been taught on the subject. His
giving the term embryo to that part of the seed which is capable of
development was also an advance in respect of logic and
morphology, in spite of his mistake in not admitting the cotyledons
which are attached to the embryo into the conception; this, however,
could easily be corrected at a later time. What Gärtner now named
the embryo, had been up to his time called the ‘corculum seminis,’
especially by Linnaeus and Jussieu; it was evidently thought that
Cesalpino’s phraseology was thus retained; but he, as we have seen,
understood by the words ‘cor seminis’ the spot where the cotyledons
spring from the germ, which spot he wrongly took for the meeting-
point of root and stem and the seat of the soul of the plant. And so
at last after two hundred years the word disappeared from use,
which might have reminded the botanist of Cesalpino’s views
respecting the soul of plants.
A work such as Gärtner’s could scarcely find a fruitful soil in
Germany, where some thirty years before even Koelreuter’s brilliant
investigations had met with little sympathy, and Conrad Sprengel’s
remarkable enquiries into the relations of the structure of the flower
to the insect-world in 1793 failed to be understood; Gärtner
complains in the second part, published in 1791, that not two
hundred copies of the first volume were sold in three years. But the
work, which forms an epoch in the history of botany, was better
received in France, where the Academy placed it as second in the list
of the productions which in later times had been most profitable to
science; there lived the man who was able to measure the whole
value of such a work—Antoine Laurent de Jussieu. But even in
Germany, where plant-describing was comfortably flourishing, there
were not altogether wanting men who knew how to estimate both
the services of Gärtner and the importance of the natural system.
First among these was August Johann Georg Karl Batsch, Professor
in Jena from 1761 to 1802, who published in the latter year a ‘Tabula
affinitatum regni vegetabilis,’ with characters of the groups and
families. Kurt Sprengel, who was born in 1766, and died as Professor
of Botany in Halle in 1833, contributed still more to the spread of
clearer views respecting the real character of the natural system and
the task of scientific botany generally by numerous works, and
especially by his ‘Geschichte der Botanik,’ which appeared in 1817
and 1818. But even this highly gifted and accomplished man agreed
with the Linnaean botanists in attributing an excessive value to the
describing of plants, as is shown in his history, where to exalt the
merits of the old botanists he gives figures of the plants first
described by them.
Meanwhile the meritorious efforts of these men were not in
themselves capable of directly advancing the natural system, or of
greatly increasing the number of its adherents in Germany, nor did it
find general acceptance in that country till it had made considerable
progress in the hands of the two foremost botanists of the time, De
Candolle and Robert Brown.

Augustin Pyrame de Candolle[37] (1778-1841) belongs to the number

of those distinguished investigators of nature, who at the end of the
last and the beginning of our own century made their native city
Geneva a brilliant centre of natural science. De Candolle was the
contemporary and fellow-countryman of Vaucher, Theodore de
Saussure, and Senebier. Physics and physiology especially were
being successfully cultivated at that time in Geneva, and Pyrame de
Candolle was attracted to these studies; among his youthful efforts
are some important investigations into the effect of light on
vegetation, and the contributions which he made to vegetable
physiology in his great work on that subject will be noticed in a later
portion of this history. De Candolle turned his attention to all parts of
theoretical and applied botany, but his importance for the history of
the science lies chiefly in the direction of morphology and systematic
botany, and it is this which we will now proceed to describe.
The amount and compass of De Candolle’s labours as a systematic
and descriptive botanist exceed those of any writer before or after
him. He wrote a series of comprehensive monographs of large
families of plants, and published a new edition of De Lamarck’s large
‘Flore Française’ substantially altered and enlarged; and in addition
to these and many similar works and treatises on the geographical
distribution of plants, he set on foot the grandest work of descriptive
botany that is as yet in existence, the ‘Prodromus Systematis
Naturalis,’ in which all known plants were to be arranged according
to his natural system and described at length,—a work not yet fully
completed, and in which many other descriptive botanists of the last
century participated, but none to so large an extent as De Candolle,
who alone completed more than a hundred families. It is not
possible to give an account in few words of the service rendered to
botany by such labours as these; they form the real empirical basis
of general botany, and the better and more carefully this is laid, the
greater the security obtained for the foundations of the whole
science.
But a still higher merit perhaps can be claimed for De Candolle,
inasmuch as he not only like Jussieu elaborated the system and its
fundamental principles in his descriptive works, but developed the
theory, the laws of natural classification, with a clearness and depth
such as no one before him had displayed. To this purpose he applied
morphological researches, which in profundity and wealth of thought
and in the fruitfulness of their results for the whole domain of
systematic botany far surpassed all that Linnaeus and Jussieu had
accomplished, and show us that while engaged in his splendid
labours in descriptive botany he had caught during his ten years’
residence in Paris the true spirit of modern investigation of nature,
as it had been developed by the French naturalists of the end of the
previous century. Scarcely a trace is to be found in De Candolle of
the scholasticism of Cesalpino and Linnaeus, which occasionally
makes its appearance even in Jussieu. For instance, he dealt with
morphology as essentially the doctrine of the symmetry of form in
plants, that is, he found the basis of morphological examination in
the relative position and numbers of the organs, disregarding their
physico-physiological properties as of no account from the
morphological point of view. He was therefore the first who
recognised the remarkable discordance between the morphological
characters of organs, which are of value for systematic purposes,
and their physiological adaptations to the conditions of life, though it
must at the same time be acknowledged, that he did not
consistently carry out this principle, but committed grave offences
against it in laying down his own system. It is a point of the highest
interest in De Candolle’s morphological speculations, that he was the
first who endeavoured to refer certain relations of number and form
to definite causes, and thus to distinguish what is primary and
important in the symmetry of plants from merely secondary
variations, as is seen in his doctrine of the abortion and adherence
of organs. In these distinctions De Candolle laid the foundation of
morphological views, which, though now modified to some extent,
do still contain the chief elements of morphology and the natural
system; but his morphological speculations were confined to the
domain of the Phanerogams, and chiefly advanced the theory of the
flower; a morphology of the Cryptogams was as little to be thought
of in the condition of microscopy before 1820, as the application of
the history of development to the establishment of morphological
theories.
De Candolle published his morphology or doctrine of symmetry
and his theory of classification together in a book which appeared
first in 1813, with the title, ‘Théorie Élémentaire de la botanique ou
exposition des principes de la classification naturelle et de l’art de
décrire et d’étudier les végétaux,’ and again in 1819 in an improved
and enlarged edition. The second edition will be the one referred to
in the further account of his views. The second chapter of the
second book concerns us most at present. After alluding to the fact,
that anatomy and physiology are concerned with the structure of the
individual organ only so far as the power to fulfil its proper function
depends on the structure, he points out that the physiological point
of view is no longer sufficient when we are engaged in comparing
the organs of different plants. Though it is true that the function of
the organs is the most important for the life and permanence of the
individual, yet we find these functions modified in the case of
homologous organs in different plants; for the natural classification
we must take into consideration only the entire system of
organisation, that is, the symmetry of the organs. All organisms of a
kingdom, he continues, have the same functions with slight
modifications; the immense amount of variation in systematically
different species depends therefore only on the way in which the
general symmetry of structure varies. This symmetry of the parts,
the discovery of which is the great object in the investigation of
nature, is nothing more than the sum total (l’ensemble) of the
positional relations of the parts. Whenever these relations
(disposition) are regulated according to the same plan, the
organisms exhibit a certain general resemblance to one another,
independently of the form of the organs in detail; when this general
resemblance is perceived, without any attempt to give any account
of it in the detail, we have what has been called habitual
relationship; but it is the task of the doctrine of symmetry to resolve
this likeness of habit into its elements, and to explain its causes.
Without this study of symmetry it may easily happen that two
different kinds of symmetry may be supposed to be alike, because
they seem outwardly alike to our senses, just as forms of crystals of
different systems may be confounded together for want of careful
examination; the chief thing is to know the plan of symmetry in
every class of plants, and the study of this is the foundation of every
theory of natural affinities. But success in this study depends on the
certainty with which organs are distinguished, and the distinguishing
them must be independent of changes of form, size, and function.
He then shows that the difficulties in the morphological comparison
of organs, or, as we should now say, in the establishing the
homology, are due to three causes; abortion, degeneration, and
adherence (adhérence). These three causes, by which the original
symmetry of a class is changed and may even be utterly obscured,
are then fully illustrated by examples.
In respect to abortion he distinguishes that which is produced by
internal causes from that which is due to accidental and external
ones; he refers especially to the abortion of two loculaments in the
fruit of the horse-chestnut and the oak, to the suppression of the
terminal bud in some shrubs by the adjoining axillary buds, and to
the fact that all organs of plants may become abortive in a similar
manner; for instance, the sexual organs disappear entirely in the
disk-flowers of Viburnum Opulus, and one of the two sexes in the
flower of Lychnis dioica. He goes on to answer the question, how it
is possible to discover the symmetry in such cases; one method he
finds supplied by monstrosities, among which there are even some
that may be regarded as a return to the original symmetry, the cases
known as peloria. Analogy or ‘induction’ is, he says, less certain, but
of much more extensive application; this is founded exclusively on
the knowledge of the relative position of organs. Armed with this,
we find that the flower of Albuca, which corresponds to a flower of
Liliaceae in everything except in having only three stamens, is to be
considered one of the Liliaceae, because it has three filaments
placed between the three stamens exactly in the position of the
three other stamens in the Liliaceae; it must be concluded therefore
that they are abortive stamens. Similar conclusions from analogy
must be carried from species to species, from organ to organ, and
the great systematists have in fact done so. In certain cases abortion
is produced by defect, in others by excess of nourishment, of which
he gives examples. An important sentence occurs in this place;
everything in nature, he says, leads us to believe that all organisms
in their inner nature are regular, and that different forms of abortion
differently combined are the cause of all irregularity; from this point
of view the smallest irregularities are important, because they lead
us to expect greater ones in nearly allied plants; and wherever in a
given system of organisation there are inequalities between organs
of the same name, the inequality will possibly reach a maximum,
that is, end by annihilating the smallest part. Thus in the Labiatae
with two stamens, it is the two which in other cases also are the
smaller, which are here completely aborted. When in Crassulaceae
there are twice as many stamens as petals, those that alternate with
the petals are larger and earlier developed, and we may therefore
expect that those which are opposite the petals may become
abortive; and therefore we may place a genus like Sedum, in which
the latter are sometimes wanting, with Crassulaceae; but we could
not do so, if we found only the stamens that are superposed upon
the petals. It occurs sometimes, he continues, that an organ is
prevented from fulfilling its function by partial abortion. In this case
it may assume another function, as the abortive leaves of the vetch
and the abortive inflorescences of the vine are employed as tendrils.
In other cases the abortive organ appears to be quite useless, as for
instance many rudimentary leaves. All such useless organs, says De
Candolle, exist only in consequence of the primitive symmetry of all
organs. Finally the abortion may be so complete that no trace of the
organ remains, of which case there are however two kinds, one
where the organ is at first perceptible and afterwards quite
disappears, as in the abortive loculaments in the fruit of the oak; in
other instances no trace is to be seen from the first of the abortive
organs, as happens with the fifth stamen of Antirrhinum.
All that has here been said might be alleged word for word in
proof of the theory of descent, but our author is an adherent of the
dogma of the constancy of species; what from his point of view he
really means by abortion is difficult to say, for the object which is
aborted is wanting. If species are constant, and therefore of
absolutely distinct origin, we must not speak of abortion; we can
only say that an organ which is present or large in one species is
small or wanting in another. In introducing the idea of abortion De
Candolle at once goes beyond the dogma of the constancy of
species, without being clear in his own mind with regard to this
important step. His proceeding shows that facts lead even a
defender of constancy against his will to theories which run counter
to that dogma. This is confirmed by his perception of the correlation
of growth, which is connected with abortion; he points to the fact
that owing to the disappearance of sexual organs in the disk-flowers
of Viburnum Opulus the corollas become larger, as do the bracts of
the abortive flowers of Salvia Horminum; similarly he regards the
disappearance of the seeds in Ananas, Banana, and the Bread-fruit
tree as the cause of the enlargement of the pericarps; it does not
escape him, that the fertile flower-stalks in Rhus Cotinus remain
naked, while an elegant pubescence forms on the barren ones; the
leaf-like expansion of the leaf-stalks of Acacia heterophylla, which do
not develop their laminae, he refers also to this correlation of
growth. He finds the most remarkable example of the kind in the
doubling of flowers, where according to his view the disappearance
of the anthers is a condition of the corolline expansion of the
filaments; in the same way sometimes the carpel is changed into a
petal through the disappearance of the stigma. Though in many of
these cases it is quite possible to conceive of the relations of cause
and effect in the reverse way, yet De Candolle’s principle of
correlation will be equally applicable.
The second cause by which the symmetry may be obliterated,
namely degeneration, asserts itself in the formation of thorns, of
threadlike prolongations of membranous expansions, and in the
production of fleshy parts, or of parts with dry membranes.
The third kind of departure from the symmetrical plan is the
adherence of parts, the theory of which he grounds first and chiefly
on the phenomena of grafting, and then passes to more difficult
cases. The close packing of the ovaries in some species of
honeysuckle, is, he says, the primary cause of their adherence. This
therefore does not depend on the plan of symmetry, but upon an
accident, which however is constant in its appearance, owing to the
specific constitution of such plants. In connection with the
phenomena of adherence he next considers the question whether a
structure composed of several parts, as for instance a compound
ovary, should be considered as originally simple and afterwards
divided into parts, or whether the converse is the true account, and
he says that we must examine each particular case and decide which
is the correct conception. Thus it may be shown that the perfoliate
leaves of honeysuckles, as well as the involucres of many
Umbelliferae, and monosepalous calyces and monopetalous corollas
are due to adherence, and he proceeds to prove that ovaries with
several loculaments and several parts have in like manner been
formed by adherence of two or more carpellary leaves, and
concludes by pointing out the systematic importance of such
considerations. Further on he takes occasion to speak of the
significance of the relative number of the parts of the flower, on
which head he says much that is good, but does not thoroughly
investigate the matter; it was not till a later time that Schimper’s
doctrine of phyllotaxis made it possible to express these relations of
number and position more precisely.
He concludes his rules for the application of his morphology to the
determination of relations of affinity with the declaration, that the
whole art of natural classification consists in discerning the plan of
symmetry, and in making abstraction of all the deviations from it
which he has described,—much in the same way as the mineralogist
seeks to discover the fundamental forms of crystals from the many
derivative forms. It is obvious that all this teaching was a great step
in advance upon the right path, that De Candolle has here given
utterance for the first time to an important principle of morphology
and systematic botany; nevertheless he did not succeed in always
consistently carrying out his own principle; he was true to himself
only in the determination of small groups of relationship; in framing
the largest divisions of the vegetable kingdom he entirely lost sight
of the rule which he had himself laid down, that the morphological
character of organs and the extent to which it can be turned to
account for systematic purposes is entirely independent of their
physiological character, and that the most important physiological
characters are just those which are of quite subordinate importance
in the determination of affinities. In spite of this strange
inconsistency, to De Candolle belongs the merit of being the first to
point emphatically to the distinction between morphological and
physiological marks, and to bring clearly to light the discordance
between morphological affinity and physiological habit; but in this
discordance lurks a problem, which could only be solved forty years
later by Darwin’s theory of selection. A genuine inductive process
alone could reveal these remarkable relations between the
morphological and physiological characters of organs. But it is at the
same time true that De Candolle could not have made this discovery,
if his predecessors had not already established a large number of
affinities. It was while he was engaged in an exact comparison of
forms already recognised as undoubtedly related to one another,
that that which he called the plan of symmetry, and which was
afterwards named a type, revealed itself to him; and as he examined
it more closely, and compared it with peculiarities of habit in
different plants formed on the same plan, he discovered certain
causes, by means of which the deviations were to be explained;
these were abortion, degeneration, and adherence. By attending to
these he succeeded in discovering affinities that had been hitherto
doubtful or unknown; this was at all events the true inductive way of
advancing the system, and whatever the earlier systematists had
effected that was really valuable had been effected virtually in the
same way, only they never arrived at a clear understanding of their
own mode of proceeding; they had followed unconsciously the
method which De Candolle clearly understood and consciously
pursued.
The majority of De Candolle’s successors were far from fully
appreciating the entire significance of his theory, its importance as a
matter of method and principle; on the contrary in the search for
affinities they continued to surrender themselves to a blind feeling
rather than to a clearly recognised method, and the same must be
said unhappily of De Candolle himself, when he was dealing with the
establishment of the large divisions of the vegetable kingdom. With
equal surprise we find him in the book before us, in which he has
developed the true method in systematic botany, expressing the
opinion that the most important physiological characters must be
employed for the primary divisions of the system, and this idea is
not improved by the fact that he ascribes to the organs physiological
characters which they do not really possess; thus he regards the
vessels as the most important organs of nutrition, which they are not
in fact, and upon this double error he builds his primary division of
the whole vegetable kingdom into vascular and cellular plants, and
then by a third mistake believes that this division coincides with the
division of plants into those which have and those which have not
cotyledons. The already established division into Monocotyledons
and Dicotyledons, which rests upon a leading and purely
morphological mark, is spoilt by De Candolle through his following
Desfontaines in ascribing to the Dicotyledons a different mode of
growth in thickness from that of the Monocotyledons, and
characterising the one as exogenous, the other as endogenous. But
this notion is utterly incorrect, as von Mohl showed twelve years
later; and if it were correct, it would still be unimportant in a
systematic point of view, because it appeals to a mark which is
morphologically of quite subordinate importance. The worst
consequence of these mistakes was, that the Vascular Cryptogams
were introduced into the same class with the Monocotyledons,—a
decided step backwards, if we compare De Candolle’s system with
that of Jussieu. In spite of these grave defects in the primary
divisions of the whole vegetable kingdom De Candolle’s system
deserved the fame which it acquired and long maintained; it had this
advantage over Jussieu’s system that in the class of Dicotyledons,
the largest division of the whole kingdom, larger subdivisions
appeared, and these served to unite families that were in many
points essentially related; the Dicotyledons were in fact divided first
of all into two artificial groups according to the presence of two floral
envelopes or one; the first and much the larger of these was again
broken up into a series of subordinate groups, which pointed in
many ways to natural affinities. That these groups, which have only
quite recently been materially altered, did to a very considerable
extent take account of natural affinities, is due to the fact that De
Candolle in framing them really followed his own rules, whereas the
superior divisions, which are artificial, owe their existence to his
disregard of them.
De Candolle declared emphatically against the old notion, that the
vegetable system answers to a linear series,—a notion which sprang
from a misunderstanding of the saying, ‘Natura non facit saltus,’—
and demonstrated its impossibility by examples; but he allowed
himself to be too much influenced by the idea which had been
thrown out by Linnaeus, and taken up by Giseke, Batsch, Bernardin
de St. Pierre, L’Heritier, Du Petit-Thouars and others, that the
vegetable kingdom might be compared as respects its grouping to a
geographical map, in which the quarters of the globe answer to the
classes, the kingdoms to the families, and so on. If the theory of
descent is to a certain degree compatible with the idea of a linear
sequence from the most imperfect to the highest forms of plants, it
is quite incompatible with the above comparison; and systematic
investigation, led astray from the right path, is in danger of ascribing
the importance of real affinities to mere resemblances of habit,
incidental analogies, by which a group of plants appears to be
connected with five or six others. In exhibiting his system on paper
De Candolle allowed the use of the linear sequence as a
convenience, for here it was not, he said, a matter of any
importance, since the true task of the science is to study the
relations of symmetry in each family and the mutual relations of
families to one another; yet in a linear presentation of the system for
didactic purposes the sequence ought not to begin with the most
simple plants, for these are the least known, but with the most
highly developed. Thus De Candolle was the means of removing
from the system the last trace of anything which harmonised with an
ascending and uninterrupted development of forms. Resting on the
doctrine of the constancy of species, and assuming that every group
of relationship is founded on a plan of symmetry round which
individual forms are grouped as crystals round their parent form, De
Candolle was quite consistent in his views. The mode of
representation came to prevail in the vegetable kingdom which De
Candolle’s contemporary, Cuvier, an equally sturdy defender of the
dogma of constancy, had introduced in the animal kingdom as the
type-theory. Thus the most splendid results obtained by induction
were united in the case of De Candolle with the barren dogma of the
constancy of species, which, as Lange wittily remarks, comes direct
from Noah’s ark, to form an intimate mixture of truth and error; nor
did De Candolle’s many adherents succeed in unravelling the coil,
though they removed the chief errors from his system and
introduced many improvements.
To these remarks may be appended a table of the main divisions
of De Candolle’s system of 1819, which so far as it is presented in
linear arrangement he calls expressly an artificial system.
 I. Vascular plants or plants with cotyledons.

1. Exogens or Dicotyledons.
a. With calyx and corolla:
Thalamiflorals (polypetalous hypogynous),
Calyciflorals (polypetalous perigynous),
Corolliflorals (gamopetalous).
b. Monochlamydeous plants (with a single floral envelope).

2. Endogens or Monocotyledons.
a. Phanerogams (true Monocotyledons),
b. Cryptogams (vascular Cryptogams including Naiadeae).

II. Cellular plants or Acotyledons.

a. With leaves (Muscineae),

b. Without leaves (Thallophytes).

The number of families, with Linnaeus 67, with A. L. de Jussieu

100, was increased by De Candolle to 161.
If the principles of comparative morphology laid down by De
Candolle were at first prevented from being rapidly disseminated in
Germany by the philosophical tendencies then reigning among its
botanists, and especially by the obscurities of Goethe’s doctrine of
metamorphosis, yet these principles and his views also on the
natural system won their way by degrees to acknowledgment and
acceptance; and after the year 1830 the study of the system was
prosecuted by the botanists of Germany, as well as by those of
England and France, as the proper object of the science. We may
even say that the impulse given by De Candolle worked more
powerfully from that time forward in Germany than in France. It may
be said too of De Candolle’s contemporary, the Englishman Robert
Brown[38] (1773-1858), whose chief labours fall in the period
between 1820 and 1840, that he, like De Candolle, was better
appreciated during that time in Germany than in any other country.
Robert Brown, who spent the five years from 1801 to 1805 in
Australia, studied the flora of that quarter of the world, and
discussed in numerous essays the botanical results of various
journeys made by other naturalists in polar regions and in the
tropics. In this way he found opportunity to leaven the ideas, which
through Humboldt’s influence had become predominant respecting
the geography of plants, with the spirit of the natural system; he
also made the morphology and systematic position of a number of
families the subject of critical investigation.
Robert Brown’s literary efforts were limited to these monographs;
he nowhere attempted to give a connected account of the principles
which he follows in them, an exposition of his morphology or a
theory of classification, nor did he frame a new system. The results
of his studies which were really fruitful and served to advance the
science are to be found in the more general remarks, which he
managed to insert quite incidentally in his monographs. In this way
he succeeded in clearing up the morphology of the flower and with it
the systematic position of some difficult families of plants, such as
the Grasses, Orchids, Asclepiads, the newly-discovered Rafflesiaceae
and others, and to throw new light at the same time on wider
portions of the system; in his considerations on the structure and
affinities of the most remarkable plants, which had been collected in
Africa by different travellers in the years immediately following 1820,
he discussed difficult and remarkable morphological relations in the
structure of the flower. He referred especially in this essay (1826) to
the relations between the numbers of the stamens and carpels, and
those of the floral envelopes in the Monocotyledons and
Dicotyledons, and showed how these typical, or as he calls them in
De Candolle’s phraseology, symmetrical relations were changed by
abortion, while he entered at the same time into a more exact
determination of the position of the aborted and of the perfect
organs, in order to discover new relations of affinity. His most
valuable work in this direction is a paper on a genus Kingia,
discovered in New Holland in 1825; the structure of the seeds in this
genus led him to seek more accurate knowledge of the unfertilised
ovule in the Phanerogams generally, and especially in the Cycads
and Conifers. In spite of the labours of Gärtner and the later
researches of Treviranus, there was still considerable obscurity
attaching to the theory of the seed, for no one had yet succeeded in
referring the position of the embryo in the ripe seed to a general
law. For this it was necessary to submit the ovule before fertilisation
to careful examination, and Robert Brown carried out this first step
to a history of development with great success; he was the first to
distinguish the integuments and the nucleus in the ovule, and the
embryo-sac in the nucleus, parts which Malpighi and Grew had
indeed observed but had not brought out with perfect clearness. The
micropyle and the hilum of the seed had not yet been properly
distinguished, but had been to some extent even confounded with
one another. Robert Brown showed that the hilum answers to the
point of attachment of the ovule, while the micropyle is a canal
formed by the integuments of the ovule and leading to the summit
of the nucleus; that in anatropous ovules the micropyle lies beside
the hilum, in orthotropous ovules opposite to it; that the embryo in
the embryo-sac (amnion) is always formed at the spot which lies
nearest the micropyle, and that the radicle of the embryo is always
turned towards the micropyle,—facts which at once established the
general rule by which to determine the position of the embryo in the
seed and in the fruit. He also gave the first correct explanation of
the endosperm as a nourishing substance formed inside the embryo-
sac after fertilisation, and more than this, he was the first to
distinguish the perisperm as a substance formed outside the
embryo-sac in the tissue of the nucleus.
In this way Robert Brown established morphological relations in
the organisation of the seed of the Monocotyledons and
Dicotyledons, which count among the most important principles of
classification in these classes; he was still more happy in being the
first to detect the peculiar structure of the flower of Conifers and
Cycads, as compared with that of other flowering plants; it was he
who perceived that what had been hitherto called a female flower in
these plants was really a naked ovule, a view which Trew of
Nüremberg had, it is true, suggested in the year 1767. He also called
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