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Logic in Computer Science Modelling and Reasoning about Systems 2nd Edition by Michael Huth, Mark Ryan ISBN 0511261586 9780511261589 download

The document is a reference for the book 'Logic in Computer Science: Modelling and Reasoning about Systems' by Michael Huth and Mark Ryan, detailing its contents and structure. It covers various topics in logic, including propositional and predicate logic, model checking, and program verification, aimed at undergraduate and graduate students. Additionally, it provides links to download the book and other related texts.

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100% found this document useful (2 votes)
20 views55 pages

Logic in Computer Science Modelling and Reasoning about Systems 2nd Edition by Michael Huth, Mark Ryan ISBN 0511261586 9780511261589 download

The document is a reference for the book 'Logic in Computer Science: Modelling and Reasoning about Systems' by Michael Huth and Mark Ryan, detailing its contents and structure. It covers various topics in logic, including propositional and predicate logic, model checking, and program verification, aimed at undergraduate and graduate students. Additionally, it provides links to download the book and other related texts.

Uploaded by

brodergyaase
Copyright
© © All Rights Reserved
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This page intentionally left blank
LOGIC IN COMPUTER SCIENCE
Modelling and Reasoning about Systems
LOGIC IN COMPUTER SCIENCE
Modelling and Reasoning about Systems

MICHAEL HUTH
Department of Computing
Imperial College London, United Kingdom

MARK RYAN
School of Computer Science
University of Birmingham, United Kingdom
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press


The Edinburgh Building, Cambridge CB2 8RU, UK
Published in the United States of America by Cambridge University Press, New York
www.cambridge.org
Information on this title: www.cambridge.org/9780521543101

© Cambridge University Press 2004

This publication is in copyright. Subject to statutory exception and to the provision of


relevant collective licensing agreements, no reproduction of any part may take place
without the written permission of Cambridge University Press.

First published in print format 2004

ISBN-13 978-0-511-26401-6 eBook (EBL)


ISBN-10 0-511-26401-1 eBook (EBL)

ISBN-13 978-0-521-54310-1 paperback


ISBN-10 0-521-54310-X paperback

Cambridge University Press has no responsibility for the persistence or accuracy of urls
for external or third-party internet websites referred to in this publication, and does not
guarantee that any content on such websites is, or will remain, accurate or appropriate.
Contents

Foreword to the first edition page ix


Preface to the second edition xi
Acknowledgements xiii
1 Propositional logic 1
1.1 Declarative sentences 2
1.2 Natural deduction 5
1.2.1 Rules for natural deduction 6
1.2.2 Derived rules 23
1.2.3 Natural deduction in summary 26
1.2.4 Provable equivalence 29
1.2.5 An aside: proof by contradiction 29
1.3 Propositional logic as a formal language 31
1.4 Semantics of propositional logic 36
1.4.1 The meaning of logical connectives 36
1.4.2 Mathematical induction 40
1.4.3 Soundness of propositional logic 45
1.4.4 Completeness of propositional logic 49
1.5 Normal forms 53
1.5.1 Semantic equivalence, satisfiability and validity 54
1.5.2 Conjunctive normal forms and validity 58
1.5.3 Horn clauses and satisfiability 65
1.6 SAT solvers 68
1.6.1 A linear solver 69
1.6.2 A cubic solver 72
1.7 Exercises 78
1.8 Bibliographic notes 91
2 Predicate logic 93
2.1 The need for a richer language 93

v
vi Contents

2.2 Predicate logic as a formal language 98


2.2.1 Terms 99
2.2.2 Formulas 100
2.2.3 Free and bound variables 102
2.2.4 Substitution 104
2.3 Proof theory of predicate logic 107
2.3.1 Natural deduction rules 107
2.3.2 Quantifier equivalences 117
2.4 Semantics of predicate logic 122
2.4.1 Models 123
2.4.2 Semantic entailment 129
2.4.3 The semantics of equality 130
2.5 Undecidability of predicate logic 131
2.6 Expressiveness of predicate logic 136
2.6.1 Existential second-order logic 139
2.6.2 Universal second-order logic 140
2.7 Micromodels of software 141
2.7.1 State machines 142
2.7.2 Alma – re-visited 146
2.7.3 A software micromodel 148
2.8 Exercises 157
2.9 Bibliographic notes 170
3 Verification by model checking 172
3.1 Motivation for verification 172
3.2 Linear-time temporal logic 175
3.2.1 Syntax of LTL 175
3.2.2 Semantics of LTL 178
3.2.3 Practical patterns of specifications 183
3.2.4 Important equivalences between LTL formulas 184
3.2.5 Adequate sets of connectives for LTL 186
3.3 Model checking: systems, tools, properties 187
3.3.1 Example: mutual exclusion 187
3.3.2 The NuSMV model checker 191
3.3.3 Running NuSMV 194
3.3.4 Mutual exclusion revisited 195
3.3.5 The ferryman 199
3.3.6 The alternating bit protocol 203
3.4 Branching-time logic 207
3.4.1 Syntax of CTL 208
Contents vii

3.4.2 Semantics of CTL 211


3.4.3 Practical patterns of specifications 215
3.4.4 Important equivalences between CTL formulas 215
3.4.5 Adequate sets of CTL connectives 216
3.5 CTL* and the expressive powers of LTL and CTL 217
3.5.1 Boolean combinations of temporal formulas in CTL 220
3.5.2 Past operators in LTL 221
3.6 Model-checking algorithms 221
3.6.1 The CTL model-checking algorithm 222
3.6.2 CTL model checking with fairness 230
3.6.3 The LTL model-checking algorithm 232
3.7 The fixed-point characterisation of CTL 238
3.7.1 Monotone functions 240
3.7.2 The correctness of SATEG 242
3.7.3 The correctness of SATEU 243
3.8 Exercises 245
3.9 Bibliographic notes 254
4 Program verification 256
4.1 Why should we specify and verify code? 257
4.2 A framework for software verification 258
4.2.1 A core programming language 259
4.2.2 Hoare triples 262
4.2.3 Partial and total correctness 265
4.2.4 Program variables and logical variables 268
4.3 Proof calculus for partial correctness 269
4.3.1 Proof rules 269
4.3.2 Proof tableaux 273
4.3.3 A case study: minimal-sum section 287
4.4 Proof calculus for total correctness 292
4.5 Programming by contract 296
4.6 Exercises 299
4.7 Bibliographic notes 304
5 Modal logics and agents 306
5.1 Modes of truth 306
5.2 Basic modal logic 307
5.2.1 Syntax 307
5.2.2 Semantics 308
5.3 Logic engineering 316
5.3.1 The stock of valid formulas 317
viii Contents

5.3.2 Important properties of the accessibility relation 320


5.3.3 Correspondence theory 322
5.3.4 Some modal logics 326
5.4 Natural deduction 328
5.5 Reasoning about knowledge in a multi-agent system 331
5.5.1 Some examples 332
5.5.2 The modal logic KT45n 335
5.5.3 Natural deduction for KT45n 339
5.5.4 Formalising the examples 342
5.6 Exercises 350
5.7 Bibliographic notes 356
6 Binary decision diagrams 358
6.1 Representing boolean functions 358
6.1.1 Propositional formulas and truth tables 359
6.1.2 Binary decision diagrams 361
6.1.3 Ordered BDDs 366
6.2 Algorithms for reduced OBDDs 372
6.2.1 The algorithm reduce 372
6.2.2 The algorithm apply 373
6.2.3 The algorithm restrict 377
6.2.4 The algorithm exists 377
6.2.5 Assessment of OBDDs 380
6.3 Symbolic model checking 382
6.3.1 Representing subsets of the set of states 383
6.3.2 Representing the transition relation 385
6.3.3 Implementing the functions pre∃ and pre∀ 387
6.3.4 Synthesising OBDDs 387
6.4 A relational mu-calculus 390
6.4.1 Syntax and semantics 390
6.4.2 Coding CTL models and specifications 393
6.5 Exercises 398
6.6 Bibliographic notes 413
Bibliography 414
Index 418
Foreword to the first edition

by
Edmund M. Clarke
FORE Systems Professor of Computer Science
Carnegie Mellon University
Pittsburgh, PA

Formal methods have finally come of age! Specification languages, theorem


provers, and model checkers are beginning to be used routinely in industry.
Mathematical logic is basic to all of these techniques. Until now textbooks
on logic for computer scientists have not kept pace with the development
of tools for hardware and software specification and verification. For exam-
ple, in spite of the success of model checking in verifying sequential circuit
designs and communication protocols, until now I did not know of a sin-
gle text, suitable for undergraduate and beginning graduate students, that
attempts to explain how this technique works. As a result, this material is
rarely taught to computer scientists and electrical engineers who will need to
use it as part of their jobs in the near future. Instead, engineers avoid using
formal methods in situations where the methods would be of genuine benefit
or complain that the concepts and notation used by the tools are compli-
cated and unnatural. This is unfortunate since the underlying mathematics
is generally quite simple, certainly no more difficult than the concepts from
mathematical analysis that every calculus student is expected to learn.
Logic in Computer Science by Huth and Ryan is an exceptional book.
I was amazed when I looked through it for the first time. In addition to
propositional and predicate logic, it has a particularly thorough treatment
of temporal logic and model checking. In fact, the book is quite remarkable
in how much of this material it is able to cover: linear and branching time
temporal logic, explicit state model checking, fairness, the basic fixpoint

ix
x Foreword to the first edition

theorems for computation tree logic (CTL), even binary decision diagrams
and symbolic model checking. Moreover, this material is presented at a level
that is accessible to undergraduate and beginning graduate students. Nu-
merous problems and examples are provided to help students master the
material in the book. Since both Huth and Ryan are active researchers in
logics of programs and program verification, they write with considerable
authority.
In summary, the material in this book is up-to-date, practical, and ele-
gantly presented. The book is a wonderful example of what a modern text
on logic for computer science should be like. I recommend it to the reader
with greatest enthusiasm and predict that the book will be an enormous
success.

(This foreword is re-printed in the second edition with its author’s permis-
sion.)
Preface to the second edition

Our motivation for (re)writing this book


One of the leitmotifs of writing the first edition of our book was the obser-
vation that most logics used in the design, specification and verification of
computer systems fundamentally deal with a satisfaction relation
Mφ
where M is some sort of situation or model of a system, and φ is a specifi-
cation, a formula of that logic, expressing what should be true in situation
M. At the heart of this set-up is that one can often specify and implement
algorithms for computing . We developed this theme for propositional,
first-order, temporal, modal, and program logics. Based on the encourag-
ing feedback received from five continents we are pleased to hereby present
the second edition of this text which means to preserve and improve on the
original intent of the first edition.

What’s new and what’s gone


Chapter 1 now discusses the design, correctness, and complexity of a SAT
solver (a marking algorithm similar to Stålmarck’s method [SS90]) for full
propositional logic.
Chapter 2 now contains basic results from model theory (Compactness
Theorem and Löwenheim–Skolem Theorem); a section on the transitive clo-
sure and the expressiveness of existential and universal second-order logic;
and a section on the use of the object modelling language Alloy and its anal-
yser for specifying and exploring under-specified first-order logic models with
respect to properties written in first-order logic with transitive closure. The
Alloy language is executable which makes such exploration interactive and
formal.

xi
xii Preface to the second edition

Chapter 3 has been completely restructured. It now begins with a discus-


sion of linear-time temporal logic; features the open-source NuSMV model-
checking tool throughout; and includes a discussion on planning problems,
more material on the expressiveness of temporal logics, and new modelling
examples.
Chapter 4 contains more material on total correctness proofs and a new
section on the programming-by-contract paradigm of verifying program cor-
rectness.
Chapters 5 and 6 have also been revised, with many small alterations and
corrections.

The interdependence of chapters and prerequisites


The book requires that students know the basics of elementary arithmetic
and naive set theoretic concepts and notation. The core material of Chap-
ter 1 (everything except Sections 1.4.3 to 1.6.2) is essential for all of the
chapters that follow. Other than that, only Chapter 6 depends on Chapter 3
and a basic understanding of the static scoping rules covered in Chapter 2 –
although one may easily cover Sections 6.1 and 6.2 without having done
Chapter 3 at all. Roughly, the interdependence diagram of chapters is

2 3 4 5

WWW page
This book is supported by a Web page, which contains a list of errata;
text files for all the program code; ancillary technical material and links;
all the figures; an interactive tutor based on multiple-choice questions;
and details of how instructors can obtain the solutions to exercises in
this book which are marked with a ∗. The URL for the book’s page
is www.cs.bham.ac.uk/research/lics/. See also www.cambridge.org/
052154310x
Acknowledgements

Many people have, directly or indirectly, assisted us in writing this book.


David Schmidt kindly provided serveral exercises for Chapter 4. Krysia
Broda has pointed out some typographical errors and she and the other
authors of [BEKV94] have allowed us to use some exercises from that book.
We have also borrowed exercises or examples from [Hod77] and [FHMV95].
Susan Eisenbach provided a first description of the Package Dependency
System that we model in Alloy in Chapter 2. Daniel Jackson make very
helpful comments on versions of that section. Zena Matilde Ariola, Josh
Hodas, Jan Komorowski, Sergey Kotov, Scott A. Smolka and Steve Vickers
have corresponded with us about this text; their comments are appreciated.
Matt Dwyer and John Hatcliff made useful comments on drafts of Chap-
ter 3. Kevin Lucas provided insightful comments on the content of Chapter
6, and notified us of numerous typographical errors in several drafts of the
book. Achim Jung read several chapters and gave useful feedback.
Additionally, a number of people read and provided useful comments on
several chapters, including Moti Ben-Ari, Graham Clark, Christian Haack,
Anthony Hook, Roberto Segala, Alan Sexton and Allen Stoughton. Numer-
ous students at Kansas State University and the University of Birmingham
have given us feedback of various kinds, which has influenced our choice and
presentation of the topics. We acknowledge Paul Taylor’s LATEX package for
proof boxes. About half a dozen anonymous referees made critical, but con-
structive, comments which helped to improve this text in various ways. In
spite of these contributions, there may still be errors in the book, and we
alone must take responsibility for those.
Added for second edition
Many people have helped improve this text by pointing out typos and
making other useful comments after the publication date. Among them,

xiii
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Fig. 153. Fig. 154.
Tuft of chætophora, Portion of chætophora
natural size. showing branching.

320. Position of œdogonium.—Œdogonium is one of the true


thread-like algæ, green in color, and the threads are divided into
distinct cells. It, along with many relatives, was once placed in the
old genus conferva. These are all now placed in the group
Confervoideæ, that is, the conferva-like algæ.
321. Relatives of œdogonium.—Many other genera are
related to œdogonium. Some consist of simple threads, and others
of branched threads. An example of the branched forms is found in
chætophora, represented in figures 153, 154. This plant grows in
quiet pools or in slow-running water. It is attached to sticks, rocks,
or to larger aquatic plants. Many threads spring from the same point
of attachment and radiate in all directions. This, together with the
branching of the threads, makes a small, compact, greenish,
rounded mass, which is held firmly together by a gelatinous
substance. The masses in this species are about the size of a small
pea, or smaller. Growth takes place in chætophora at the ends of the
threads and branches. That is, growth is apical. This, together with
the branched threads and the tendency to form cell masses, is a
great advance of the vegetative condition of the plant upon that
which we find in the simple threads of œdogonium.
CHAPTER XVII.
COLEOCHÆTE.
322. Among the green algæ coleochæte is one of the most interesting. Several species are known
in this country. One of these at least should be examined if it is possible to obtain it. It occurs in the
water of fresh lakes and ponds, attached to aquatic plants.

Fig. 155. Fig. 156.


Stem of aquatic plant Thallus of Coleochæte scutata.
showing coleochæte,
natural size.

323. The shield-shaped coleochæte.—This plant (C. scutata) is in the form of a flattened,
circular, green plate, as shown in fig. 156. It is attached near the center on one side to rushes and other
plants, and has been found quite abundantly for several years in the waters of Cayuga Lake at its
southern extremity. As will be seen it consists of a single layer of green cells which radiate from the
center in branched rows to the outside, the cells lying so close together as to form a continuous plate.
The plant started its growth from a single cell at the central point, and grew at the margin in all
directions. Sometimes they are quite irregular in outline, when they lie quite closely side by side and
interfere with one another by pressure. If the surface is examined carefully there will be found long
hairs, the base of which is enclosed in a narrow sheath. It is from this character that the genus takes its
name of coleochæte (sheathed hair).
Fig. 157.
Portion of thallus of Coleochæte scutata,
showing empty cells from which zoogonidia
have escaped, one from each cell;
zoogonidia at the left. (After Pringsheim.) Fig. 158.
Portion of thallus of Coleochæte
scutata, showing four antheridia
formed from one thallus cell; a
single spermatozoid at the right.
(After Pringsheim.)

324. Fruiting stage of coleochæte.—It is possible at some seasons of the year to find rounded
masses of cells situated near the margin of this green disk. These have developed from a fertilized egg
which remained attached to the plant, and probably by this time the parent plant has lost its color.
325. Zoospore stage.—This mass of tissue does not develop directly into the circular green disk,
but each of the cells forms a zoospore. Here then, as in œdogonium, we have another stage of the
plant interpolated between the fertilized egg and that stage of the plant which bears the gametes. But
in coleochæte we have a distinct advance in this stage upon what is present in œdogonium, for in
coleochæte the fertilized egg develops first into a several-celled mass of tissue before the zoospores are
formed, while in œdogonium only four zoospores are formed directly from the egg.
326. Asexual reproduction.—In asexual reproduction any of the green cells on the plant may
form zoogonida. The contents of a cell round off and form a single zoogonidium which has two cilia at
the smaller end of the oval body, fig. 157. After swimming around for a time they come to rest,
germinate, and produce another plant.
327. Sexual reproduction.—Oogonium.—The oogonium is formed by the enlargement of a cell
at the end of one of the threads, and then the end of the cell elongates into a slender tube which opens
at the end to form a channel through which the spermatozoid may pass down to the egg. The egg is
formed of the contents of the cell (fig. 159). Several oogonia are formed on one plant, and in such a
plant as C. scutata they are formed in a ring near the margin of the disk.
Fig. 159.
Coleochæte soluta; at left branch bearing oogonium (oog); antheridia (ant);
egg in oogonium and surrounded by enveloping threads; at center three antheridia
open, and one spermatozoid; at right sporocarp, mature egg inside sporocarp wall.

Fig. 161.
Fig. 160. Sporocarp ruptured by growth
Two sporocarps of egg to form cell mass.
still surrounded Cells of this sporophyte
by thallus. forming zoospores.
Thallus finally
decays and sets
sporocarp free.

Figs. 160, 161. C. scutata.

328. Antheridia.—In C. scutata certain of the cells of the plant divide into four smaller cells, and
each one of these becomes an antheridium. In C. soluta the antheridia grow out from the end of
terminal cells in the form of short flasks, sometimes four in number or less (fig. 159). A single
spermatozoid is formed from the contents. It is oval and possesses two long cilia. After swimming
around it passes down the tube of the oogonium and fertilizes the egg.
329. Sporocarp.—After the egg is fertilized the cells of the threads near the egg grow up around it
and form a firm covering one cell in thickness. This envelope becomes brown and hard, and serves to
protect the egg. This is the “fruit” of the coleochæte, and is sometimes called a sporocarp (spore-fruit).
The development of the cell mass and the zoospores from the egg has been described above.
Some of the species of coleochæte consist of branched threads, while others form circular cushions
several layers in thickness. These forms together with the form of our plant C. scutata make an
interesting series of transitional forms from filamentous structures to an expanded plant body formed of
a mass of cells.

330. COMPARATIVE TABLE FOR SPIROGYRA, VAUCHERIA, ŒDOGONIUM, COLEOCHÆTE.

GAMETOPHYTE. (Bears the sexual organs and gonidia.)


SPORO
Vegetative Mulitipl- Sexual Reproduction. Bears
Growth.
Phase ication.
Sexual Organs. Gametes. Fr
Simple Undifferentiated. Undifferentiated.
All cells By
threads of Zyg
Spirogyra. divide and breaking up Any cell of thread. Entire contents of
cylindrical Rests.
grow. of threads. Conjugate by tube. cojugating cells.
cells.
Differentiated. Differentiated.
By
Oogonium,
multiciliate
Limited large rounded
Branched zoogonidia, Antheridia Egg
to ends of cell on special Small two- Large
Vaucheria. threads, and other slender cells oospo
threads and branch, opens ciliated egg
continuous. cells, from on special Rests.
branches. and emits bit spermatozoids. cell.
terminal branches.
of
portions.
protoplasm.
Differentiated. Differentiated.
By oval Antheridia
Oogonium,
zoogonidia, disk-shaped, Oval
Simple Limited changed
with crown several from spermatozoids Egg
threads of to certain vegetative Large
Œdogonium. of cilia. Any one with crown of oospo
cylindrical portions of cell, opens egg
cell may vegetative cilia. Two from Rests.
cells. thread. and emits bit cell.
form a single cell. each
of
zoogonidium. Sometimes on antheridium.
protoplasm.
dwarf males.
Differentiated. Differentiated.
Oogonium,
enlarged veg.
cell, with long
Egg
By tube through
(surro
Branched zoogonidia opening of
Antheridia, Oval, by wa
threads, or Terminal with two which
four or biciliate Large gamet
Coleochæte. compact or cilia. Any cell spermatozoid
several from spermatozoid, egg Rests.
circular marginal. may form a enters. After
single veg. one from each cell. and g
plates. single fertilization
cell. ntheridium. form
zoogonidium. wall of
of cells
enveloping
threads
surrounds
oogonium.
CHAPTER XVIII.
CLASSIFICATION AND ADDITIONAL
STUDIES OF THE ALGÆ.
In order to show the general relationship of the algæ studied, the
principal classes are here enumerated as well as some of the
families. In some of the groups not represented by the examples
studied above, a few species are described which may serve as the
basis of additional studies if desired. The principal classes[17] of
algæ are as follows:
Class Chlorophyceæ.
331. These are the green algæ, so called because the
chlorophyll green is usually not masked by other pigments, though in
some forms it is. There are three subclasses.
332. Subclass PROTOCOCCOIDEÆ.—In the Protococcoideæ
are found the simplest green plants. Many of them consist of single
cells which live an independent life. Others form “colonies,” loose
aggregations of individuals not yet having attained the permanency
of even a simple plant body, for the cells often separate readily and
are able to form new colonies. The colonies are often held together
by a gelatinous membrane, or matrix. Some are motile, while others
are non-motile. A few of the families are here enumerated.
333. Family Volvocaceæ.—These are all motile, during the
vegetative stage. The individuals are single or form more or less
globose colonies.
334. The “red snow” plant (Sphærella nivalis).—This is
often found in arctic and alpine regions forming a red covering over
more or less large areas of snow or ice. For this reason it is called
the “red snow plant.”
335. Sphærella lacustris, a closely related species, is very
widely distributed in temperate regions along streams or on the
borders of lakes and ponds. Here in dry weather it is often found
closely adhering to the dry rock surface, and giving it a reddish color
as if the rock were painted. This is especially the case in the shallow
basins formed over the uneven surface of the rock near the water’s
edge. These places during heavy rains or in high water are provided
with sufficient water to fill the basins. During such times the red
snow plant grows and multiplies, loses its red color and becomes
green, and, being motile, is free swimming. It is a single-celled
plant, oval in form, surrounded by a gelatinous sheath and with two
cilia or flagella at the smaller end, by the vibration of which it moves
(fig. 162). The single cell multiplies by dividing into two cells. When
the water dries out of the basin, the motile plant comes to rest, and
many of the cells assume the red color. To obtain the plant for study,
scrape some of the red covering from these rock basins and place it
in fresh spring water, and in a day or so the swarmers are likely to
be found. Under certain conditions small microzoids are formed.

Fig. 162.
Sphærella lacustris (Girod.) Wittrock. A, mature
free swimming individual with central red spot. B,
division of mother individual to form two. C, division of
a red one to form four. D, division into eight. E, a
typical resting cell, red. F, same beginning to divide. G,
one of four daughter zoospores after swimming around
for a time losing its red color and becoming green.
(After Hazen.)
336. Chlamydomonas is a very interesting genus of motile
one-celled green algæ, because the species are closely related to the
Flagellates among the lower animals. The plant is oval, with a single
chloroplast and surrounded by a gelatinous envelope through which
the two cilia or flagella extend. One-celled organisms of this kind are
sometimes called monads, i.e., a one-celled being. This one has a
gelatinous cloak and is, therefore, a cloaked monad
(Chlamydomonas). The species often are found as a very thin green
film on fresh water. C. pulvisculus is shown in fig. 163. When it
multiplies the single cell divides into two, as shown in B. Sometimes
a non-motile palmella stage is formed, as shown in C and D.
Reproduction takes place by gametes which are of unequal size, the
smaller one representing the sperm and the larger one the egg, as
in E and F. These conjugate as in G and H, the protoplasm of the
smaller one passing over into the larger one, and a zygospore is thus
formed.

Fig. 163.
Chlamydomonas pulvisculus (Müll.) Ehrb. A, an old
motile individual; n, nucleus; p, pyrenoid; s, red eye
spot; v, contractile vacuole; B, motile individual has
drawn in its cilia and divided into two; C, mother plant
has drawn in its cilia and divided into four non-motile
cells; D, pamella stage; E, female gamete—egg; F,
male gamete—sperm; G, early stage of conjugation;
H, zygospore with conjugating tube and empty male
cell attached. (After Wille.)
337. Of those which form colonies, Pandorina morum is
widely distributed and not rare. It consists of a sphere formed of
sixteen individuals enclosed in a thin gelatinous membrane. Each cell
possesses two cilia (or flagella), which extend from the broader end
out through the enveloping membrane. By the movement of these
flagella the colony goes rolling around in the water. When the plant
multiplies each individual cell divides into sixteen small cells, which
then grow and form new colonies. Reproduction takes place when
the individual cells of the young colonies separate, and usually a
small individual unites with a larger one and a zygospore is formed
(see fig. 164). Eudorina elegans is somewhat similar, but when the
gametes are formed certain mother cells divide into sixteen small
motile males or sperms, and certain other mother cells divide into
sixteen large motile females or eggs. These separate from the
colonies, and the sperms pair with the eggs and fuse to form
zygospores. This plant as well as Chlamydomonas pulvisculus
foreshadows the early differentiation of sex in plants.
Fig. 164.
Pandorina morum (Müll.) Bory.
I, motile colony;
II, colony divided into
16 daughter colonies;
III, sexual colony, gametes escaping;
IV, V, conjugating gametes;
VI, VII, young and old zygospore;
VIII, zygospore forming a large swarm
spore, which is free in IX;
X, same large swarm spore divided
to form young colony.
(After Pringsheim.)
Fig. 165.
Pleurococcus
(protococcus)
vulgaris.

338. Family Tetrasporaceæ.—This family is well represented


by Tetraspora lubrica forming slimy green net-like sheets attached to
objects in slow-running water. It is really a single-celled plant. The
rounded cells divide by cross walls into four cells, and these again,
and so on, large numbers being held in loose sheets by the slime in
which they are imbedded.
339. Family Pleurococcaceæ.—The members of this family
are all non-motile in the vegetative stage. They consist of single
individuals, or of colonies. Pleurococcus vulgaris (Protococcus
vulgaris) is a single-celled alga, usually obtained with little difficulty.
It is often found on the shaded, and cool, or moist side of trees,
rocks, walls, etc., in damp places. This plant is not motile. It
multiplies by fission (Fig. 165) into two, then four, etc. These cells
remain united for a time, then separate. Sometimes the cells are
found growing out into filaments, and it is thought by some that P.
vulgaris may be only a simple stage of a higher alga. Eremosphæra
viridis is another single-celled alga found in fresh water among
filamentous forms. The cells are large and globose.

Fig. 166.
Pediastrum boryanum. A, mature colony, most of
the young colonies have escaped from their mother
cells; at g, a young colony is escaping; sp, empty
mother cells; B, young colony; C, same colony with
spores arranged in order. (After Braun.)
340. Family Hydrodictyaceæ.—These plants form colonies of
cells. Hydrodictyon reticulatum, the water net, is made up of large
numbers of cylindrical cells so joined at their ends as to form a large
open mesh or net. Pediastrum forms circular flat colonies, as shown
in fig. 166. Both of these plants are rather common in fresh-water
pools, the latter one intermingled with filamentous algæ, while the
former forms large sheets or nets. Multiplication in Hydrodictyon
takes place by the protoplasm in one of the cells dividing into
thousands of minute cells, which gradually arrange themselves in the
form of a net, escape together from the mother cell, and grow into a
large net. In Pediastrum multiplication takes place in a similar way,
but the protoplasm in each cell usually divides into sixteen small
cells, and escaping together from the mother cell arrange
themselves and grow to full size (fig. 166).
341. The Conjugateæ include several families of green algæ,
which probably should be included among the Chlorophyceæ. They
have probably had their origin from some of the more simple
members of the Protococcoideæ. They are represented by Spirogyra,
Zygnema, and the desmids, studied in Chapter 14.
342. Subclass CONFERVOIDEÆ.—These are mostly
filamentous algæ, the filaments being composed of cells firmly
united, and, with the exception of the simplest forms, there is a
definite growing point. A few of the families are as follows:
343. Family Ulvaceæ.—These contain the sea wracks, or sea
lettuce, like Ulva, forming expanded green, ribbon-like growths in
the sea.
Fig. 167.
Ulothrix zonata. A, base of thread. B, cells with
zoospores, C, one cell with zoospores escaping another
cell with small biciliate gametes escaping and some
fusing to form zygospores, E, zoospores germinating
and forming threads: F, G, zygospore growing and
forming zoospores. (After Caldwell and Dodel-Port.)
344. Family Ulotrichaceæ, represented by Ulothrix zonata, not
uncommon in slow-running water or in ponds of fresh water
attached to rocks or wood. It consists of simple threads of short
cells. Multiplication takes place by zoospores. Reproduction takes
place by motile sexual cells (gametes) which fuse to form a
zygospore (fig. 167).
345. Family Chætophoraceæ, represented by Chætophora (in
Chapter 15) and Drapernaudia in fresh water.
346. Family Œdogoniaceæ, represented by Œdogonium
(Chapter 16).
347. Family Coleochætaceæ, represented by Coleochæte
(Chapter 17).
348. Subclass SIPHONEÆ.—There are several families.
349. Family Botrydiaceæ.—This is represented by Botrydium
granulatum (Chapter 15, p. 146).
350. Family Vaucheriaceæ, represented by Vaucheria
(Chapter 15), with quite a large number of species, is widely
distributed.
Class Schizophyceæ (= Cyanophyceæ).
351. The Blue-Green Algæ, or
Cyanophyceæ form slimy looking thin
mats on damp wood or the ground, or
floating mats or scum on the water. The
color is usually bluish green, but in some
species it is purple, red or brown. All have
chlorophyll, but it is not in distinct
chloroplasts and is more or less completely
guised by the presence of other pigments.
Two orders and eight families are
recognized. The following include some of Fig. 168.
our common forms: Glœocapsa.
352. ORDER COCCOGONALES
(COCCOGONEÆ).—Single-celled plants,
occurring singly or in colonies, in some forms forming short threads.
One of the two families is mentioned.
353. Family Chroococcaceæ.—The plants multiply only
through cell division. Chroococcus, forms rounded, blue-green cells
enclosed in a thick gelatinous coat, in fresh water and in damp
places; certain species form “lichen-gonidia” in some genera of
lichens. Glœocapsa is similar to Chroococcus, but the colonies are
surrounded by an additional common gelatinous envelope (fig. 168);
on damp rocks, etc.
Fig. 169.
A, Oscillatoria princeps, a, terminal cell; b, c,
portions from the middle of a filament. In c, a dead
cell is shown between the living cells; B, Oscillatoria
froelichii, b, with granules along the partition walls.

354. ORDER HORMOGONALES (HORMOGONEÆ).—Plants


filamentous, simple celled or with false or true branching, usually
several celled (Spirulina is single celled). Multiplication takes place
through hormogones, short sections of the threads becoming free;
also through resting cells. Two of the six families are mentioned.
355. Family Oscillatoriaceæ.—This family is represented by
the genus Oscillatoria, and by several other genera common and
widely distributed. Oscillatoria contains many species. They are
found on the damp ground or wood, or floating in mats in the water.
They often form on the soil at the bottom of the pool, and as gas
becomes entangled in the mat of threads, it is lifted from the bottom
and floated to the surface of the water. The plant is thread-like, and
divided up into many short cells. The threads often show an
oscillating movement, whence the name Oscillatoria.
356. Family Nostocaceæ.—This family is represented by
Nostoc, which forms rounded, slimy, blue-green masses on wet
rocks. The individual plants in the slimy ball resemble strings of
beads, each cell being rounded, and several of these arranged in
chains as shown in fig. 170. Here and there are often found larger
cells (heterocysts) in the chain. Nostoc punctiforme lives in the
intercellular spaces of the roots of cycads (often found in
greenhouses), and in the stems of Gunnera. N. sphæricum lives in
the spaces between the cells in many species of liverworts (in the
genera Anthoceros, Blasia, Pellia, Aneura, Riccia, etc.), and in the
perforated cells of Sphagnum acutifolium. Anabæna is another
common and widely distributed genus. The species occur in fresh or
salt water, singly or in slimy masses. Anabæna azollæ lives
endophytically in the leaves of the water fern, Azolla.

Fig. 170.
Nostoc linckii. A, filament with two heterocysts (h),
and a large number of spores (sp); B, isolated spore
beginning to germinate; C, young filament developed
from spore. (After Bornet.)

Fig. 171.
Bacteria. A, Bacillus subtilis. Spores in threads,
unstained rods, and stained rods showing cilia; B,
Bacillus tetani, the tetanus or lockjaw bacillus, found in
garden soil and on old rusty nails. Spores in club-
shaped ends. C, Micrococcus; D, Sarcina; E,
Streptococcus; F, Spirillum. (After Migula.)
Class Schizomycetes.
357. Bacteriales.—The bacteria are sometimes classified with
the Cyanophyceæ, under the name Schizophyta, and represent the
subdivision Schizomycetes, or fission fungi, because many of them
multiply by a division of the cells just as the blue-green algæ do. For
example, Bacillus forms rods which increase in length and divide into
two rods, or it may grow into a long thread of many short rods.
Micrococcus consists of single rounded cells. Streptococcus forms
chains of rounded cells, Sarcina forms irregular cubes of rounded
cells, while others like Spirillum are spiral in form. Bacillus subtilis
may be obtained by making an infusion from hay and allowing it to
stand for several days. Bacillus tetani occurs in the soil, on old rusty
nails, etc. It is called the tetanus bacillus because it causes a
permanent spasm of certain muscles, as in “lockjaw.” This bacillus
grows and produces this result on the muscles when it occurs in
deep and closed wounds such as are caused by stepping on an old
nail or other object which pierces the flesh deeply. In such a deep
wound oxygen is deficient, and in this condition the bacillus is
virulent. Opening the wounds to admit oxygen and washing them
out with a solution of bichloride of mercury prevents the tetanus.
Many bacteria are of great importance in bringing about the decay of
dead animal and plant matter, returning it to a condition for plant
food. (See also nitrate and nitrite bacteria, Chapter IX.) While most
bacteria are harmless there are many which cause very serious
diseases of man and animals, as typhoid fever, diphtheria,
tuberculosis, etc., while some others produce disease in plants.
Others aid in certain fermentations of liquids and are employed for
making certain kinds of wines or other beverages. Some work in
symbiosis with yeasts, as in the kephir yeast, used in fermenting
certain crude beverages by natives of some countries.
357a. Myxobacteriales (Myxobacteriaceæ Thaxter[18]).—
These plants consist of colonies of bacteria-like organisms, motile
rods, which multiply by cross-division and secrete a gelatinous
substance or matrix which surrounds the colonies. They form
plasmodium-like masses which superficially resemble the slime
moulds. In the fruiting stage some species become elevated from
the substratum into cylindrical, clavate, or branched forms, which
bear cysts of various shapes containing the rods in a resting stage,
or the rods are converted into spore-like masses. Ex., Chondromyces
crocatus on decaying plant parts, Myxobacter aureus on wet wood
and bark, Myxococcus rubescens on dung, decaying lichens, paper,
etc.
Class Flagellata.
358. The flagellates are organisms of very low organization
resembling animals as much as they do plants. They are single
celled and possess two cilia or flagella, by the vibration of which
they move. Some are without a cell wall, while others have a well-
defined membrane, but it rarely consists of cellulose. Some have
chromatophores and are able to manufacture carbohydrates like
ordinary green plants. These are green in Euglena, and brown in
Hydrurus. Some possess a mouth-like opening and are able to ingest
solid particles of food (more like animals), while others have no such
opening and absorb food substances dissolved in water (more like
plants). The Euglena viridis is not uncommon in stagnant water,
often forming a greenish film on the water.
Class Peridineæ.
358a. These are peculiar one-celled organisms provided with two
flagella and show some relationship to the Flagellates. They usually
are provided with a cellulose membrane, which in some forms
consists of curiously sculptured plates. In the higher forms this
cellulose membrane consists of two valves fitting together in such a
way as to resemble some of the diatoms. Like the Flagellates, some
have green chromatophores, which in some are obscured by a
yellow or brown pigment (resembling the diatoms), while still others
have no chlorophyll. The Peridineæ are abundant in the sea, while
some are found in fresh water.
Class Diatomaphyceæ
(Bacillariales, Diatomaceæ).

Fig. 171a.
A group of Diatoms: c and d, top and side views of
the same form; e, colony of stalked forms attached to
an alga; f and g, top and side views of the form shown
at e; h, a colony; i, a colony, the top and side view
shown at k and n, forming auxospores. (After Kerner.)
358b. The diatoms are minute and peculiar organisms believed
to be algæ. They live in fresh, brackish, and salt water. They are
often found covering the surface of rocks, sticks, or the soil in thin
sheets. They occur singly and free, or several individuals may be
joined into long threads, or other species may be attached to objects
by slender gelatinous stalks. Each protoplast is enclosed in a silicified
skeleton in the form of a box with two halves, often shaped like an
old-fashioned pill box, one half fitting over the other like the lid of a
box. It is evident that in this condition the plant cannot increase
much in size.
They multiply by fission. This takes place longitudinally, i.e., in
the direction of the two halves or valves of the box. Each new plant
then has a valve only on one side. A new valve is now formed over
the naked half, and fits inside the old valve. At each division the
individuals thus become smaller and smaller until they reach a
certain point, when the valves are cast off and the cell forms an
auxospore, i.e., it grows alone, or after conjugation with another, to
the full size again, and eventually provides itself with new valves.
The valves are often marked, with numerous and fine lines, often
making beautiful figures, and some are used for test objects for
microscopes.
The free forms are capable of movement. The movement takes
place in the longitudinal direction of the valves. They glide for some
time in one direction, and then stop and move back again. It is not a
difficult thing to mount them in fresh water and observe this
movement.
The diatoms have small chlorophyll plates, but the green color is
disguised by a brownish pigment called diatomin. The relationships
of the diatoms are uncertain, but some, because of the color, think
they are related to the Phæophyceæ.
Class Phæophyceæ.
359. The brown algæ. (Phæophyceæ).—The members of
this class possess chlorophyll, but it is obscured by a brown pigment.
The plants are accessible at the seashore, and for inland laboratories
may be preserved in formalin (2½ per cent). (See also Chapter LVI.)

Fig. 172.
A, Ectocarpus siliculosus; B, branch with a young
and a ripe plurilocular sporangium; E, gametes fusing
to form zygospore, (B, after Thuret; E, after Berthold.)
360. Ectocarpus.—The genus Ectocarpus represents well some
of the simpler forms of the brown algæ (fig. 172). They are slender,
filamentous branched algæ growing in tufts, either epiphytic on
other marine algæ (often on Fucaceæ), or on stones. The slender
threads are only divided crosswise, and thus consist of long series of
short cells. The sporangia are usually plurilocular (sometimes
unilocular), and usually occur in the place of lateral branches. The
zoospores escape from the apex of the sporangium and are biciliate,
and they fuse to form zygospores.
361. Sphacelaria.—The species of this genus represent an
advance in the development of the thallus. While they are
filamentous and branched, division takes place longitudinally as well
as crosswise (fig. 173).
362. Leathesia difformis represents an interesting type
because the plant body is small, globose, later irregular and hollow,
and consists of short radiately arranged branches, the surface ones
in the form of short, crowded, but free, trichome-like green
branches. This trichothallic body recalls the similar form of
Chætophora pisiformis (Chapter 16) among the Chlorophyceæ.
Fig. 173.
Sphacelaria, portion of plant
showing longitudinal division
of cells, and brood bud
(plurilocular sporangium).
Fig. 174.
Laminaria digitata, forma cloustoni,
North Sea. (Reduced.)

363. The Giant Kelps.—Among the brown algæ are found the
largest specimens, some of the laminarias or giant kelps, rivaling in
size the largest land plants, and some of them have highly
developed tissues. Postelsia palmæformis has a long, stout stem,
from the free end of which extend numerous large and long blades,
while the stem is attached to the rocks by numerous “root” like
outgrowths, the holdfasts. It occurs along the northern Pacific coast,
and appears to flourish where it receives the shock of the surf
beating on the shore. Several species of Laminaria occur on our
north Atlantic coast. In L. digitata, the stem expands at the end into
a broad blade, which becomes split into several smaller blades (fig.
174). Macrocystis pyrifera inhabits the ocean in the southern
hemisphere, and sometimes is found along the north American
coast. It is said to reach a length of 200-300 meters.
364. Fucus, or Rockweed.—This plant is a more or less
branched and flattened thallus or “frond.” One of them, illustrated in
fig. 119, measures 15-30 cm (6-12 inches) in length. It is attached
to rocks and stones which are more or less exposed at low tide.
From the base of the plant are developed several short and more or
less branched expansions called “holdfasts,” which, as their name
implies, are organs of attachment. Some species (F. vesiculosus)
have vesicular swellings in the thallus.
The fruiting portions are somewhat thickened as shown in the
figure. Within these portions are numerous oval cavities opening by
a circular pore, which gives a punctate appearance to these fruiting
cushions. Tufts of hairs frequently project through them.
Fig. 175.
Portion of plant of Fucus
showing conceptacles in
enlarged ends; and below
the vesicles
(Fucus vesiculosus).
Fig. 176.
Section of conceptacle
of Fucus, showing oogonia,
and tufts of antheridia.

365. Structure of the conceptacles.—On making sections of


the fruiting portions one finds the walls of the cavities covered with
outgrowths. Some of these are short branches which bear a large
rounded terminal sac, the oogonium, at maturity containing eight
egg-cells. More slender and much-branched threads bear narrowly
oval antheridia. In these are developed several two-ciliated
spermatozoids.
Fig. 177.
Oogonium of Fucus
with ripe eggs.
Fig. 178.
Antheridia of Fucus,
on branched threads.

366. Fertilization.—At maturity the spermatozoids and egg-


cells float outside of the oval cavities, where fertilization takes place.
The spermatozoid sinks into the protoplasm of the egg-cell, makes
its way to the nucleus of the egg, and fuses with it as shown in fig.
181. The fertilized egg then grows into a new plant. Nearly all the
brown algæ are marine.
Fig. 180.
Fig. 179. Eggs of Fucus surrounded
Antheridia of Fucus with by spermatozoids.
escaping spermatozoids.

Fig. 181.
Fertilization in Fucus; fn, female nucleus; mn, male
nucleus; n, nucleolus. In the left figure the male
nucleus is shown moving down through the cytoplasm
of the egg; in the remaining figures the cytoplasm of
the egg is omitted. (After Strasburger.)
367. The Gulf weed (Sargassum bacciferum) in the warmer
Atlantic ocean unites in great masses which float on the water,
whence comes the name “Sargassum Sea.” The Sargassum grows on
the coast where it is attached to the rocks, but the beating of the
waves breaks many specimens loose and these float out into the
more quiet waters, where they continue to grow and multiply
vegetatively.
368. Uses.—Laminaria japonica and L. angustata are used as
food by the Chinese and Japanese. Some species of the
Laminariaceæ are used as food for cattle and are also used for
fertilizers, while L. digitata is sometimes employed in surgery.
Classification.—Kjellman divides the Phæophyceæ into two
orders.
369. Order Phæosporales (Phæosporeæ) including 18
families. One of the most conspicuous families is the Laminariaceæ,
including among others the Giant Kelps mentioned above (Laminaria,
Postelsia, Macrocystis, etc.).
370. Order Cyclosporales (Cyclosporeæ).—This includes one
family, the Fucaceæ with Ectocarpus, Sphacelaria, Læathesia, Fucus,
Sargassum, etc.
Class Rhodophyceæ.
371. The red algæ (Rhodophyceæ).—The larger number of
the so-called red algæ occur in salt water, though a few genera
occur in fresh water. The plants possess chlorophyll, but it is usually
obscured by a reddish or purple pigment.
372. Nemalion.—This is one of the lower marine forms, though
its thallus is not one of the simplest in structure. The plant body
consists of a slender cylindrical branched shoot, sometimes very
profusely branched. The central strand is rather firm, while the
cortex is composed of rather loose filaments.
Fig. 182.
A red alga (Nemalion). A, sexual branches, showing
antheridia (a); carpogonium or procarp (o) with its
trichogyne (t), to which are attached two spermatia
(s); B, beginning of a cystocarp (o), the trichogyne (t)
still showing; C, an almost mature cystocarp (o), with
the disorganizing trichogyne (t). (After Vines.)
373. Batrachospermum.—This genus occurs in fresh water,
and the species are found in slow-running water of shallow streams
or ditches. There is a central slender strand which is more or less
branched, and on these branches are whorls of densely crowded
slender branches occurring at regular intervals. The plants are
usually very slippery. Gonidia are formed on the ends of some of
these branches in globose sporangia, called monosporangia, since
but a single spore or gonidium is developed in each. Other branches
often terminate in long slender hyaline setæ.
374. Lemanea.—This genus also occurs in fresh water. The
species develop only during the cold winter months in rapids of
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