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An Introduction to Plant Structure
and Development
Plant Anatomy for the Twenty-First Century
Second Edition

This is a plant anatomy textbook unlike any other on the market today.
As suggested by the subtitle, it is plant anatomy for the twenty-first cen-
tury. Whereas traditional plant anatomy texts include primarily descriptive
aspects of structure with some emphasis on patterns of development, this
book not only provides a comprehensive coverage of plant structure, but
also introduces, in some detail, aspects of the mechanisms of development,
especially the genetic and hormonal controls, and the roles of the cytoskele-
ton. The evolution of plant structure and the relationship between structure
and function are also discussed throughout the book. Consequently, it pro-
vides students and, perhaps, some teachers as well, with an introduction
to many of the exciting, contemporary areas at the forefront of research,
especially those areas concerning development of plant structure. Those
who wish to delve more deeply into areas of plant development will find
the extensive bibliographies at the end of each chapter indispensible. If
this book stimulates a few students to become leaders in teaching and
research in plant anatomy of the future, the goal of the author will have
been accomplished.

charles b. beck, Professor Emeritus of Botany at the University of Michi-

gan, received his PhD degree from Cornell University where he developed
an intense interest in the structure of fossil and living plants under the
influence of Professor Harlan Banks and Professor Arthur Eames. Following
post-doctoral study with Professor John Walton at Glasgow University in
Scotland, he joined the faculty of the University of Michigan. At Michigan
he served one term each as Chairman of the Department of Botany and
Director of the Museum of Paleontology. His graduate students pursued
research in either plant structure and development or paleobotany. He
taught courses in plant anatomy, plant morphology and paleobotany over
a period of 35 years.
An Introduction to
Plant Structure and
Development
Plant Anatomy for the
Twenty-First Century
Second Edition
Charles B. Beck
University of Michigan
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore,
São Paulo, Delhi, Dubai, Tokyo

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/9780521518055
© Charles B. Beck

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 2010

ISBN-13 978-0-511-77022-7 eBook (NetLibrary)

ISBN-13 978-0-521-51805-5 Hardback

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.
 To
My wife, Janice,
and our daughters, Ann and Sara
for their love, encouragement,
and enduring support,
and
to my students,
David Benzing
Robert Chau
Crispin Devadas
Margaret Knaus
G. Kadambari Kumari
Rudolf Schmid
William Stein
Garland Upchurch
Richard White
David Wight

who are a continuing inspiration and

from whom I have learned much.
It is important that students bring
a certain ragamuffin barefoot
irreverence to their studies; they
are not here to worship what is
known but to question it.
Jacob Bronowski
The Ascent of Man (1975)
 Contents

Preface to the second edition page xiii

Preface xv
Acknowledgements xvii
General references xx

Chapter 1 Problems of adaptation to a terrestrial

environment 1
Perspective: the origin of vascular plants 1
Structural adaptations 3
Preview of subsequent chapters 5
References 6
Further reading 7

Chapter 2 An overview of plant structure

and development 8
Perspective: origin of multicellularity 8
Some aspects of the shoot system of the vascular plant 10
Apical meristems 13
Primary tissue regions of the stem and root 17
Vascular bundle types 23
Secondary growth 26
Cells of the xylem 29
Cells of the phloem 33
References 35
Further reading 36

Chapter 3 The protoplast of the eukaryotic cell 38

Perspective 38
Morphology of the protoplast 39
Vacuoles 51
Movement of organelles in the protoplast 51
Ergastic substances 53
References 55
Further reading 57

Chapter 4 Structure and development of the cell wall 58

Perspective 58
Structure and composition of the cell wall 59
Growth of the cell wall 65
Cell wall development 68
viii CONTENTS

Plasmodesmata 72
References 77
Further reading 81

Chapter 5 Meristems of the shoot and their role in

plant growth and development 83
Perspective 83
Apical meristems 83
Formation of leaf primordia 90
Transitional tissue regions 92
Intercalary meristems 95
The primary peripheral thickening meristem of
monocotyledons 96
Cell growth and development 97
The effect of hormones on cell growth and development 98
Genetic control of cell growth and development 99
Role of the cytoskeleton in cell growth and development 100
Cell shaping by microtubules 102
References 103
Further reading 106

Chapter 6 Morphology and development of the

primary vascular system of the stem 108
Perspective 108
Cellular composition and patterns of development of primary
xylem 109
Cellular composition and patterns of development of primary
phloem 112
Differentiation of primary vascular tissues 113
The role of auxin in the development of the primary vascular
system 119
References 121
Further reading 122

Chapter 7 Sympodial systems and patterns

of nodal anatomy 123
Perspective: leaf traces 123
Nodal structure of pteridophytes 123
Sympodial systems of seed plants 125
Leaf trace lacunae 134
The cauline vs. foliar nature of vascular bundles in the eustele 135
Phyllotaxy 136
References 139
Further reading 140
 CONTENTS ix

Chapter 8 The epidermis 141

Perspective 141
Epidermis of the shoot 141
Epidermis of the root 149
Stomata 149
Signal transduction and movement in guard cells 152
Development of stomata 153
References 155
Further reading 156

Chapter 9 The origin of secondary tissue systems and

the effect of their formation on the primary
body in seed plants 157
Perspective: role of the vascular cambium 157
The effect of secondary growth on the primary body 160
The effect of secondary growth on leaf and branch traces 162
References 165
Further reading 165

Chapter 10 The vascular cambium: structure

and function 166
Perspective 166
Structure of the vascular cambium 166
General overview of cambial activity 169
Plant hormones and cambial activity 173
Submicroscopic structure of cambial initials 174
The onset of dormancy and the reactivation of dormant
cambium 175
Cytokinesis in fusiform initials 177
The problem of differential growth of cambial cells and
immature cambial derivatives 178
References 179
Further reading 182

Chapter 11 Secondary xylem 184

Perspective 184
Overview of the structure of secondary xylem 184
Secondary xylem of gymnosperms 187
Resin ducts 193
Secondary xylem of dicotyledons 194
Differentiation of tracheary elements 199
Patterns of distribution of xylary elements and rays 207
Tyloses 211
Evolution in secondary xylem of dicotyledons 213
Mechanism of water transport 215
x CONTENTS

References 216
Further reading 220

Chapter 12 The phloem 222

Perspective: evolution of the phloem 222
Gross structure and development of the phloem 223
The nature and development of the cell wall of sieve elements 228
Role of the cytoskeleton in wall development 231
The nature and development of the protoplast of sieve elements 232
Nature and function of P-protein 234
Distinctive features of the phloem of gymnosperms 236
The nature and function of companion cells and Strasburger
cells 237
The mechanism of transport in the phloem 241
References 242
Further reading 245

Chapter 13 Periderm, rhytidome, and the nature

of bark 247
Perspective 247
Periderm: structure and development 247
Formation of rhytidome 250
Lenticels 252
The outer protective layer of monocotyledons 253
References 254
Further reading 254

Chapter 14 Unusual features of structure and

development in stems and roots 255
Perspective 255
Primary peripheral thickening meristem 255
Secondary growth in monocotyledons 256
Anomalous stem and root structure 257
References 263
Further reading 263

Chapter 15 Secretion in plants 264

Perspective 264
Substances secreted by plants 264
Mechanisms of secretion 265
Internal secretory structures 266
External secretory structures 270
References 276
Further reading 278
 CONTENTS xi

Chapter 16 The root 279

Perspective: evolution of the root 279
Gross morphology 281
Contractile roots and other highly specialized root systems 282
Apical meristems 283
The quiescent center and its role in development 286
Primary tissues and tissue regions 288
Lateral transport of water and minerals in the young root 295
Development of primary tissues 297
Auxin and tissue patterning 301
Lateral root development 302
Adventitious roots 306
Secondary growth 306
The root cap: its function and role in gravitropism 307
Mycorrhizae 312
Nitrogen fixation in root nodules 313
Root–stem transition 315
References 316
Further reading 321

Chapter 17 The leaf 324

Perspective: evolution of the leaf 324
Basic leaf structure 325
Leaf development 334
The role of the cytoskeleton in leaf development 340
The role of genetics in leaf development 341
Variations in leaf form, structure, and arrangement 343
Structure in relation to function 346
Photosynthesis and phloem loading 346
Leaf structure of C3 and C4 plants 348
Supporting structures in leaves 349
Transfusion tissue in conifers 350
Leaf abscission 351
References 353
Further reading 357

Chapter 18 Reproduction and the origin

of the sporophyte 361
Perspective: the plant life cycle 361
Reproduction in gymnosperms 362
Reproduction in angiosperms 366
Development of the seed in angiosperms 373
Fruit development and the role of fruits in seed dispersal 377
Seed germination and development of the seedling 380
Floral morphogenesis 381
Pollen–pistil interactions 384
xii CONTENTS

Self-incompatibility 387
Role of the cytoskeleton in pollen tube growth 388
References 390
Further reading 394

Glossary 398
Index 433
 Preface to the second edition

Although it has been only five years since this book was first published,
research activity during this period in many areas of plant develop-
ment has resulted in much new and important information. The basic
information on plant structure is quite stable. As a result, inclusion of
new information about various aspects of development comprise the
major changes in this 2nd edition. In addition, a new section on the
evolution of tracheary elements has been added.
The areas expanded and/or upgraded include the structure and func-
tion of the cytoskeleton, and its roles in cell wall formation and pollen
tube tip growth; the role of auxin and other hormones in development,
especially in the development of tracheary elements, as well as in cam-
bial activity and tissue patterning, and the role of PIN proteins in the
movement of auxin from cell to cell by auxin efflux transporters. The
discussion on the mechanism of movement of stomatal guard cells has
been expanded and improved. Sections on long-distance transport in
the secondary xylem and phloem have been modified to emphasize
widely accepted mechanisms of transport, and the discussion of bidi-
rectional transport in the phloem has been expanded. The discussion
of gravitropism has been brought up to date. Finally, throughout the
book, discussions of the role of genetics in plant development have
been expanded.
I believe the changes listed above have made the book more useful to
advanced students and researchers without adversely affecting its use-
fulness as an introductory plant anatomy textbook. It is not designed to
be used as the only source of information in a course in plant anatomy,
i.e., to take the place of the teacher, but rather, to be a supplement to
the teacher’s lectures and a means for the student to reinforce infor-
mation from the teacher and the laboratory exercises. The book can, of
course, also be an original source of information for students beyond
that provided by the teacher. When used in a course that emphasizes
development, the student will have the opportunity to expand his or
her knowledge of plant structure, and in a course that emphasizes
plant structure, the student can expand his or her knowledge of plant
development. Ideally, however, a twenty-first century course in plant
anatomy should consist of an integration of structure and develop-
ment. It is this ideal that I have tried to promote in the design and
preparation of this book.
For granting me permission to use photographs in this 2nd edition
of the book, I express my appreciation to university and commercial
publishers and all other copyright holders. I am especially grateful to
Professor Philip Gingerich for allowing me to use the facilities and
services of the University of Michigan Museum of Paleontology, and to
three very talented persons in this unit, Bonnie Miljour, senior scientific
 Another Random Document on
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 FOOTNOTES:
[1] Danish: Edderkop.

TRANSCRIBER’S NOTES:
Obvious typographical errors have been corrected.
Inconsistencies in hyphenation have been standardized.
Archaic or alternate spelling has been retained from the original.
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