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Plant Patterning: Structural and Molecular Genetic Patterning... Structural and Molecular Genetic Aspects ESRA GALUN The Weizmann Institute of Science, Israel Library of Congress Cataloging-in-Publication

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  • PLANT PATTERNING Structural and Molecular Genetic Aspects

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  • N E W J E R S E Y • L O N D O N • S I N G A P O R E • B E I J I N G • S H A N G H A I • H O N G K O N G • TA I P E I • C H E N N A I

    World Scientific


    PLANT PATTERNING Structural and Molecular Genetic Aspects

    ESRA GALUN The Weizmann Institute of Science, Israel

  • Library of Congress Cataloging-in-Publication Data Galun, Esra, 1927-

    Plant patterning : structural and molecular genetic aspects / Esra Galun. p. cm.

    Includes bibliographical references and index. ISBN-13: 978-981-270-408-5 (hardcover : alk. paper) ISBN-10: 981-270-408-6 (hardcover : alk. paper) 1. Plant pattern formation. I. Title.

    QK731.G32 2007 567.918--dc22


    British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library.

    For photocopying of material in this volume, please pay a copying fee through the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, USA. In this case permission to photocopy is not required from the publisher.

    Typeset by Stallion Press Email:

    All rights reserved. This book, or parts thereof, may not be reproduced in any form or by any means, electronic or mechanical, including photocopying, recording or any information storage and retrieval system now known or to be invented, without written permission from the Publisher.

    Copyright © 2007 by World Scientific Publishing Co. Pte. Ltd.

    Published by

    World Scientific Publishing Co. Pte. Ltd.

    5 Toh Tuck Link, Singapore 596224

    USA office: 27 Warren Street, Suite 401-402, Hackensack, NJ 07601

    UK office: 57 Shelton Street, Covent Garden, London WC2H 9HE

    Printed in Singapore.

    SC - Plant patterning.pmd 9/20/2007, 11:22 AM1

  • This book is dedicated to my grandchildren,

    Michal, Reuth, Guy, Nir, Ran, Shai, Chen and Dor


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    Every entity has Roots and Tips of branches And every entity has a Beginning and an End The one who is putting his mind to the Former and to the Latter He is Approaching the WAY!

    (From the DA XUE – The Great Learning, one of the Four Books of the Chinese cultural heritage)

    “Alle Theile der Pflanze – Stengel und Wurzel ausgenommen – können auf die Form des Blattes zurückgeführt werden; sie sind nichts anderes als Modifikationen derselben.”

    All parts of the plant – except the shoot and the root – can be attributed to the structure of the leaf; they are nothing but modifications of leaves.

    (From the Doctoral Thesis of Caspar Friedrich Wolff, submitted in 1759 to the University of Halle, Germany, in Latin; the German version is of 1812). Places and times may have strange associations: shortly after this German version was published in Leipzig there was the great defeat of Napoleon at the “Battle of the Nations” ... in Leipzig).


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  • These two mottos represent two extremely different lines of thought. In the Chinese motto, the tips of the branches and roots were used as metaphors. It was probably intended to bring up two issues. One is that the entities should be studied thoroughly, from the roots to branch tips. The other issue is that a thorough knowledge can lead a person to the WAY. The Way has several meanings. Kung fu-Tze (Confucius) meant by Way morality and value. The Way can also mean all the events happening between the first step and the final goal.

    In the German motto, from Caspar Friedrich Wolff, the plant is not a metaphor. It serves as an example of an approach to understand the development of organisms. Wolff was a pioneer in this approach, although he was ignored in the recent literature on plant develop- ment. This literature cites the claim of Goethe that floral members are leaves that underwent metamorphosis. Goethe’s claim was made sev- eral years after the statement of Wolff.

    This book aims to serve to bridge the gap between the metaphoric approach of Da Xue and the “down-to-earth” approach of Wolff. I shall attempt to update a wide range of readers on what we know about the processes that regulate the formation of patterns in plants. I shall also indicate clearly what is not yet known. By “plants” I mean angiosperm plants. Other, lower plants, will be mentioned only rarely. The deliberations will be on different levels of complexity, such as single cells, meristems, tissues and plant organs (e.g. roots, shoots, floral members). Wherever applicable the use of methodolo- gies such as genetics, molecular biology and structural analyses will be described, and I shall attempt to show how the application of these methodologies led to additional knowledge on the control of plant patterning.

    As with my previous books (Galun and Breiman, 1997; Galun and Galun, 2001; Galun, 2003; Galun, 2005a) the present book is aimed at a wide range of readers. Basic knowledge of plant biology, genetics and molecular biology should be helpful for the readers. However, where required, the various studies on patterning and the method- ologies that were used in these studies, will be introduced and explained. Thus, the book should be of interest and useful for the novice in molecular developmental biology as well.

    viii Plant Patterning: Structural and Molecular Genetic Aspects

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  • Because the book is comprehensive with respect to its topics (dealing with different plant components), scholars who are engaged in the study of specific aspects of plant differentiation, may benefit from the overall view of the present book. On the other hand, this book should also be useful as a companion-text for courses on plant development.

    I am grateful to my wife, Professor Margalith Galun, and to my son, Professor Eithan Galun, who read parts of the manuscript and provided useful remarks. I am also grateful to the staff members of my Plant Sciences Department for useful discussion, among them Professor Dan Atsmon and Professor Moshe Feldman, and especially Dr. Yuval Eshed, who provided very useful advice throughout the whole process of writing this book. I thank professor Dr. Rudolf Hagemann, of the University of Halle (Germany), for the German motto from thesis of C. F. Wolff. Thanks are due to Mrs. Suzanne Trauffer Ramakrishna for her professional typing. I very much appre- ciate the efforts of my publisher, World Scientific Publishing Co., espe- cially of Ms. Sook Cheng Lim, scientific editor at WSPC, for the editing and production of this book. I acknowledge the professional work of the Graphic Arts Department of the Weizmann Institute of Science for performing very professional work on the figures and on the cover. I further acknowledge the permissions to use figures and tables that were granted by the respective publishers and authors. I pro- vided the sources of these figures and tables below each figure or table and the full citations are listed in the list of references. These citations should mean that I am grateful for the consents of the publishers to use their copyright material in this book. My special gratitude goes to Dr. Nillie Weinstein. Dr. Weinstein showed great interest in my writing throughout my last four books and since then has become an inseparable component of my writings. She encouraged me to go on writing this book. I and the book benefitted immensely from her wisdom and knowledge, especially, but not only, in philosophy.

    Esra Galun Rehovot, Israel 7th April 2007

    Preface ix

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    Preface vii

    Chapter 1 Introduction 1

    Part A Basic Issues Relevant to Plant Patterning 27

    Chapter 2 Plant Hormones 29 Chapter 3 The Cytoskeleton of Plant Cells 87 Chapter 4 Phylogeny, Taxonomy and Gene Duplication 105 Chapter 5 Regulation of Gene Expression 127 Chapter 6 Systems Biology 145

    Part B The Patterning of Plant Components 157

    Chapter 7 Patterning of the Angiosperm Embryo 159 Chapter 8 The Shoot Apical Meristem 181 Chapter 9 Patterning of Roots 223 Chapter 10 The Vegetative Shoot and Transition 275

    to Flowering Chapter 11 The Angiosperm Leaf 331 Chapter 12 Patterning of Flowers and their Members 383

    Epilogue 441

    References 443

    Corpus Acknowledgments 487

    Index 489


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  • 1


    Chapter 1


    Biological Patterning: From Awareness to Understanding

    The attitude of humans towards patterns of plants and animals has gone through several transitions during the cultural evolution of man. First, there was primarily a bewilderment of nature, and this bewil- derment concerned primarily the physical world and less the biological nature. Later came an epoch during which the developmental stages that result in mature animals and plants were followed and described. Only subsequently were experimental procedures employed i

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