i Essentials of Developmental Plant Anatomy ii iii ii jj Essentials of Developmental Plant Anatomy Taylor A. Steeves Vipen K. Sawhney ii jj 1 iv 1 Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America. © Oxford University Press 2017 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. CIP data is on file at the Library of Congress ISBN 978–0 –1 9–0 65705–5 9 8 7 6 5 4 3 2 1 Printed by Sheridan Books, Inc., United States of America v Dedicated to the memory of the Late Taylor Steeves (co- author of this book). A distinguished scholar, teacher, gentleman, mentor, and dear friend. VKS vi vii ii Contents Preface ix 1. Introduction 1 2. The Plant Cell 7 The Protoplast 8 The Cell Wall 13 Cell Growth 16 3. The Flower 17 Inflorescence Types 19 Flower Morphology 19 Variations in Flower Morphology 22 The Induction of Flowering 27 Flower Development 29 4. Reproduction 31 Vegetative Reproduction 32 Sexual Reproduction 34 Overview of Sexual Reproduction 42 Apomixis 43 5. Embryo, Seed, and Fruit Development 44 Patterns of Embryo Development 44 Somatic Embryogenesis 51 Seed Development 52 Fruit Development 55 vii viii viii i Contents 6. Shoot Morphology and Development 57 Growth of the Shoot 59 Shoot Apex and the Shoot Apical Meristem 60 Shoot Expansion 63 Shoot Branching 65 Shoot Modifications 66 Reproductive Shoots 67 7. Plant Cells and Tissues 69 Cell Types and Tissues 70 8. Tissues of the Stem 81 The Dicotyledonous Stem 82 Three-Dimensional Organization of the Vascular System 84 Differentiation of Tissues in the Stem 86 The Monocotyledonous Stem 91 The Stele 92 9. The Leaf 95 Leaf Form 96 Tissues of the Leaf 101 Variations Related to the Environment 104 Leaf Development 106 10. The Root 111 Root Systems 112 Root Associations 113 Root Apex and the Root Apical Meristem 115 Tissues of the Root 118 Root Branching 123 Shoot Buds from Roots 125 11. The Secondary Body 127 The Vascular Cambium 128 Secondary Xylem (The Wood) 133 Secondary Phloem 136 Secondary Growth in Monocotyledons 138 Periderm 139 Glossary 143 Bibliography 155 Index 157 ix Preface ii The seed to put together this book was sown several years ago by the late Professor Taylor Steeves and myself with the objective of providing, for a grow- ing community of students and researchers in plant developmental genetics and molecular biology, an abridged version of plant anatomy. At the time of our proposal to Oxford University Press, there were excellent plant anatomy texts available, including The Anatomy of Seed Plants by Katherine Esau, Plant Anatomy by Abraham Fahn, Plant Anatomy (Parts 1 and 2) by Elizabeth Cutter, and Plant Anatomy by James Mauseth, and more recently other books such as Esau’s Plant Anatomy by Ray Evert and Plant Anatomy: An Applied Approach by David F. Cutler, Ted Botha, and Dennis Wm. Stevenson. Although most of these books in plant anatomy are superb in their content, they are exhaustive in detail. By using sev- eral examples from different groups of plants, they provide a comparative account of plant structure and function as well as evolutionary trends in plant structure. We felt then, and even now, a need for an abbreviated text in anatomy with a plant developmental perspective. This book should be useful to researchers and students in plant biology in general, and in developmental genetics and molec- ular biology in particular. This book could also serve as a complementary text to an excellent laboratory exercises volume, Teaching Plant Anatomy by R. Larry Peterson, Carol Peterson, and Lewis H. Melville, published in 2008. It has taken a long time to bring this project to fruition, and there are sev- eral reasons for this. Initially, Prof. Steeves and I tried to fit the writing work in ix