Encyclopedia of

Plant Physiology New Series Volume 10

Editors

A. Pirson, G6ttingen M.H. Zimmermann, Harvard

Hormonal Regulation of Development II The Functions of Hormones from the Level of the Cell to the Whole Plant

Edited by Tom K Scott

Contributors

M.L. Evans W. P. Jacobs H. Kaldewey A.C. Leopold A.G. Matthysse A.W. Naylor L.D. Nooden lA. Romberger T.K Scott T.l Wodzicki S. ZajStczkowski

With 42 Figures

Springer-Verlag Berlin Heidelberg New York Tokyo 1984

Editor: Professor TOM K. SCOTT Department of Biology The University of North Carolina Chapel Hill, North Carolina 27514/USA

Planning Volumes 9-11 : Professor N. P. KEFFORD College of Tropical Agriculture and Human Resources University of Hawaii at Manoa Honolulu, Hawaii 96822/USA

ISBN-13: 978-3-642-67733-5 e-ISBN-13: 978-3-642-67731-1 DOl: 10.1007/978-3-642-67731-1

Library of Congress Cataloging in Publication Data. Main entry under title: The functions of honnones from the level of the cell to the whole plant. (Hormonal regulation of development; 2) (Encyclopedia of plant physiology; new ser., v. 10). Includes indexes. 1. Plant hormones. I. Scott, Tom K. II. Evans, M.L. III. Series. IV. Series: Encyclopedia of plant physiology; new ser., v. 10. QK711.2.E5 vol. 10 581.1s [581.1 '4)83-27185 [QK731).

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks.

Under § 54 of the Gennan Copyright Law where copies are made for other than private use, a fee is payable to "Verwertungsgesellschaft Wort" Munich.

© by Springer-Verlag Berlin· Heidelberg 1984

Softcover reprint of the hardcover 1st edition 1984

The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.

2131/3130-543210

In memory of

Martin H. Zimmermann whose dedication and enthusiasm for Developmental Botany was an inspiration to all who knew him

T.K. Scott

In Memoriam

With the death of Martin Huldrych Zimmermann on March 7, 1984, at the age of 58, the international community of plant physiology has lost not only a colleague whose own practical scientific work has played a decisive role in the development of this field, but also the author and editor of numerous scien­tific works that have greatly served the dissemination of scientific knowledge.

We mourn for a friend, who as long ago as 1974 entrusted a book to us, (Zimmermann and Brown: "Trees "). On the recent founding of a new book series (" Springer Series in Wood Science "), Martin H. Zimmermann willingly contributed his own work as the initial volume ("Xylem Structure and the Ascent of Sap "). This was to be his last book; the moving epilog - placed at his own wish at the last moment at the conclusion of the volume - became also his farewell to active publishing.

Martin H. Zimmermann's main work in the long years of cooperation with Springer-Verlag was, however, the handbook series "Encyclopedia of Plant Physiology, New Series". Over more than 12 years of intensive and often labori­ous effort he achieved, together with his editorial colleague, the realization of a complete survey of plant physiology. All those who experienced personally with what patience, efficiency, and diplomacy Martin H. Zimmermann not only planned each volume, but also indefatigably supported the numerous editors and authors with good counsel and help, will feel, with us, a great debt of gratitude to him.

The first volume of this series was edited by him jointly with 1.A. Milburn. Tragically, it was no longer granted to him to see the series brought to its conclusion.

Springer-Verlag Spring 1984

ANDRE PIRSON

Foreword

This is the second of the set of three volumes in the Encyclopedia of Plant Physiology, New Series, that will cover the area of the hormonal regulation of plant growth and development. The overall plan for the set assumes that this area of plant physiology is sufficiently mature for a review of current knowl­edge to be organized in terms of unifying principles and processes. Reviews in the past have generally treated each class of hormone individually, but this set of volumes is subdivided according to the properties common to all classes. Such an organization permits the examination of the hypothesis that differing classes of hormones, acting according to common principles, are determinants of processes and phases in plant development. Also in keeping with this theme, a plant hormone is defined as a compound with the properties held in common by the native members of the recognized classes of hormone.

Current knowledge of the hormonal regulation of plant development is grouped so that the three volumes consider advancing levels of organizational complexity, viz: molecular and subcellular; cells, tissues, organs, and the plant as an organized whole; and the plant in relation to its environment.

The first volume, entitled Molecular Aspects of Plant Hormones and edited by Professor J. MACMILLAN, was published in 1980. It treated the molecular and subcellular aspects of hormones and the processes they regulate. Although it dealt with chemically distinct classes of hormone, the first volume stressed properties and modes of studying them that were common to all classes.

In the present volume, the roles of hormones at levels of organization from the cell up to the whole plant are traced. The cellular processes of increase and change, and the interrelations of cells in tissues, of tissues in organs, and of organs in the whole plant, are considered in turn. During this progressive treatment of levels of organization, the relevant basic properties of hormones are introduced and illustrated.

A third volume addresses the interrelationships of hormones with factors in the environments of the tissues, the organs and the whole plants, within which the hormones are functioning. When this volume touches upon wide­reaching topics such as photomorphogenesis or plant movements, only those aspects that relate to principles of hormonal regulation are treated. Separate volumes of the Encyclopedia of Plant Physiology, New Series, provide compre­hensive treatment of topics such as photomorphogenesis and plant movements.

My role in the preparation of these volumes has been to propose a theme and prepare a plan to cover the current status of the field of hormonal regula­tion, then to circumscribe the portions of the plan that form logical volumes. Thereafter, the editors of the individual volumes have determined the manner in which the domain for which they accepted responsibility was treated. The

x Foreword

editor of the present volume is Professor TOM K. SCOTT and, in the Introduction to his volume, he outlines his approach and that of his authors.

The base from which these volumes have developed is the old series of the Encyclopedia of Plant Physiology. The volumes in the New Series of the Encyclopedia may, therefore, concentrate on principles that may be derived from the mass of older information and on the findings of the past 20 years. The length of each volume has been deliberately restricted, but effective organi­zation of topics and their succinct treatment assures the reader of a concise but comprehensive statement of current knowledge and thought in the field.

I thank Professor Kenneth V. Thimann for reviewing the theme and initial plan for these volumes with me.

May 1984 N.P. KEFFoRD

Contents

Introduction

T.K. SCOTT.

1 Hormonal Regulatory Systems in Plants A.C. LEOPOLD and L.D. NOODEN

1.1 Introduction ....... . . . . . . . . . . . . . . . . 1.1.1 The Concept of the Plant Hormone as a Chemical Messenger 1.1.2 Types of Hormones ............. .

1.2 Hormone Messenger Systems . . . . . . . . . . . . . 1.2.1 Common Difficulties with Single Hormonal Controls 1.2.2 Is a Hormone Always a Hormone? . . . . . . 1.2.3 Further Complications in the Hormone Concept 1.2.4 Regulatory Sites in Hormonal Controls 1.2.5 Patterns in Multiple Hormonal Control 1.2.6 Complications Arising from Compartmentation

1.3 Analysis of Hormonal Controls . . . . . 1.3.1 Two Major Approaches from the Past 1.3.2 Analyses of Hormone Systems

1.4 Conclusions ............ . 1.4.1 The Complexity of Hormonal Controls 1.4.2 Nonhormonal Correlative Controls . . 1.4.3 Evolutionary Plasticity of Hormonal Controls 1.4.4 The Future

References . . . . . . . . . . . . . . . . . . . .

2 Functions of Hormones at the Cellular Level of Organization M.L. EVANS (With 1 Figure)

2.1 Introduction .. . . . . . . . . . . . . . . . 2.2 Promotive and Inhibitory Classes of Plant Hormones 2.3 Modes of Hormone Interaction . . . . . . . . .

2.3.1 Interaction and Independence in Hormone Action 2.3.2 Synergism and Antagonism in Hormone Action

2.3.2.1 Synergism ............ . 2.3.2.2 Antagonism ........... .

2.3.3 Hormone Effects on Endogenous Hormone Levels 2.3.3.1 Hormona: Modification of Endogenous Auxin 2.3.3.2 Hormonal Modification of Endogenous Gibberellin 2.3.3.3 Hormonal Modification of Endogenous Ethylene 2.3.3.4 Hormonal Modification of Endogenous Abscisic Acid 2.3.3.5 Hormonal Modification of Endogenous Cytokinin . . 2.3.3.6 Summary .................. .

2.3.4 Influence of Environmental Parameters on Endogenous Hormones 2.3.4.1 Light 2.3.4.2 Water Stress

4 4 5 6 6 7 8 9

10 11 11 12 13 15 15 15 16 16 17

23 23 24 24 25 25 25 27 27 28 29 29 30 30 31 31 33

XII

2.3.4.3 Temperature . . . . . 2.3.4.4 Mechanical Stimulation 2.3.4.5 Summary

Contents

2.4 Hormonal Regulation . . . . .

34 34 34 35 35 36 39 40 42 43 44 44 46 48 48 48 49 49 50 51 52 53 54 54 55 55 56 56 56 56 57 57 57 57 57 59 61 62

2.4.1 The Cell Cycle . . . . . . 2.4.1.1 Effect of Cytokinins 2.4.1.2 Effect of Auxins . . 2.4.1.3 Effects of Other Growth Regulators

2.4.2 Endomitosis 2.4.3 Meiosis . . . . . . . . . . . . . . 2.4.4 Summary . . . . . . . . . . . . .

2.5 Hormone Effects on the Plane of Cell Division 2.6 Relationship of Cell Multiplication to Cell Expansion 2.7 Hormonal Regulation of Cell Expansion

2.7.1 Stem Cells ....... . 2.7.1.1 Effects of Auxin . . . . . 2.7.1.2 Effects of Gibberellin 2.7.1.3 Effects of Cytokinin, Ethylene, and Abscisic Acid

2.7.2 Root Cells ....... . 2.7.2.1 Effects of Auxin . . . 2.7.2.2 Effects of Gibberellin 2.7.2.3 Effects of Ethylene 2.7.2.4 Effects of Abscisic Acid 2.7.2.5 Effects of Cytokinin

2.7.3 Leaf Cells . . . . . . . . 2.7.3.1 Effects of Auxin . . 2.7.3.2 Effects of Gibberellin 2.7.3.3 Effects of Ethylene 2.7.3.4 Effects of Abscisic Acid 2.7.3.5 Effects of Cytokinin

2.7.4 Other Cell Types . . . 2.7.4.1 Pollen Tubes 2.7.4.2 Epidermal Hairs

2.7.5 Lower Plants 2.7.5.1 Algae 2.7.5.2 Fungi

2.7.6 Summary References . . . . . .

3 Transport and Other Modes of Movement of Hormones (Mainly Auxins) H. KALDEWEY (With 7 Figures)

3.1 Introduction . . . . . . . . . . . . . . . . . 80 3.2 Historical Sketch . . . . . . . . . . . 80

3.2.1 Indirect Evidence for Hormone Translocation 80 3.2.2 Direct Evidence for Hormone Translocation 81 3.2.3 Hormone Transport Experiments . . . . . . 81

3.2.3.1 The Early Period ......... 81 3.2.3.2 The Period of Confirmation, Extension, and Criticism 82 3.2.3.3 The" Modern" Period . . . . . 83

3.3 General Remarks on Hormone Movement .......... 86 3.3.1 Terminology and the Transport Equation . . . . . . . . 86 3.3.2 Methods to Estimate Hormone Translocation Characteristics 88

3.3.2.1 Intercept Method of van der Weij 88 3.3.2.2 Short-Term Collecting Method 92 3.3.2.3 Pulse Experiments . . . . . . . 94

Contents XIII

3.3.2.4 Penetration-Depletion Method ............. 98 3.3.2.5 Electric Potential Wave Method . . . . . . . . . . . . . 101

3.3.3 Evaluation of Estimation and Interpretation of Hormone Movement 103 3.3.3.1 Does Auxin Move in a Stream? . 103 3.3.3.2 Loading of the Transport System 105 3.3.3.3 Validity of Velocity Estimations . 106 3.3.3.4 Fluctuations of Transport Velocity 108 3.3.3.5 Flexibility of the Transport System 109

3.3.4 Models for Transport Mechanisms 111 3.3.4.1 Conveyor Belt Model 112 3.3.4.2 Electrical Polarity Hypothesis 112 3.3.4.3 Polar Secretion Theory . . . 112 3.3.4.4 Theory of Chemiosmotic Polar Diffusion 115

3.3.5 Binding Sites ........ 116 3.3.6 Pathways of Hormone Movement 117

3.3.6.1 Shoots . . . . . . . 117 3.3.6.2 Roots . . . . . . . . 118

3.4 Evidence of Hormone Translocation . 119 3.4.1 Hormone Movement in Vascular Systems 119 3.4.2 Hormone Translocation in Shoot Sections 122

3.4.2.1 Auxins . . . . . . . . . . . . 122 3.4.2.2 Hormones Other than Auxins . . 123

3.4.3 Hormone Translocation in Root Sections 125 3.4.3.1 Auxins . . . . . . . . . . 125 3.4.3.2 Hormones Other than Auxins 126

References . . . . . . . . . . . . . . . . 126

4 Functions of Hormones at Tissue Level of Organization W.P. JACOBS (With 12 Figures)

4.1 Introduction .. . . . . . . . . . . . . . . . . . 149 4.2 Role of Hormones in Differentiation of Vascular Tissues 149

4.2.1 Regeneration in Shoots . . 149 4.2.1.1 Xylem . . . . . . . . . . . . . . . . 149 4.2.1.2 Sieve-Tube Elements . . . . . . . . . . 154 4.2.1.3 Fibers . . . . . . . . . . . . . . . . 159

4.2.2 Differentiation During Normal Development in Shoots 159 4.2.2.1 Xylem . . . . . . 159 4.2.2.2 Sieve-Tube Elements 160 4.2.2.3 Fibers . . . . . . 160 4.2.2.4 Cambium 162

4.2.3 Control of Differentiation in Roots 165 4.2.4 Evidence of Differentiation from Tissue Cultures 165

4.3 General Discussion 166 References . . . . . . . . . . . . . . . . . . . . . 168

5 Functions of Hormones at the Organ Level of Organization A.W. NAYLOR (With 8 Figures)

5.1 Introduction. . . . . . . . . . . . . . . 172 5.2 Embryogeny - Establishment of Polarity 174 5.3 Hormonal Relations of the Developing Embryo 177 5.4 Vascular Devclopment- Spatial Arrangement 178 5.5 Regeneration 180 5.6 Flower Initiation . . 186 5.7 Flower Development 192

XIV

5.8 Fruit Setting and Development . . . . . 5.9 Seed Development . . . . . . . . . . 5.10 Seed Dormancy- Germination Inhibitors 5.11 Conclusions References . . . . . . . . . . . . . . . .

6 Functions of Hormones at the Whole Plant Level of Organization A.G. MATTHYSSE and T.K. SCOTT (With 8 Figures)

6.1 Introduction " . . . . . . . . . . . . . . . . . . . 6.2 Some Perspectives on the Nature of Hormones ..... . 6.3 A General View of the Role of Hormones in the Whole Plant 6.4 Some Considerations Regarding Experimental Design 6.5 Some Considerations Regarding Research Strategies 6.6 Conclusion References . . . . . . . . . . . . . . . . . . . .

7 Auxin Waves and Plant Morphogenesis S. ZAJ~CZKOWSKI, T.J. WODZICKI, and J.A. ROMBERGER (With 6 Figures)

7.1 Introduction . . . . . . . . . . . . . . . . . 7.2 Polar Transport of Auxin and Positional Information 7.3 The Origin of a New Concept 7.4 Some Experimental Evidence 7.5 The Proposed Model 7.6 Further Implications References . . . . . .

Author Index

Subject Index

Contents

195 200 204 207 208

219 220 221 224 224 226 226

244 244 245 246 252 258 259

263

289

List of Contributors

M.L. EVANS Department of Botany Ohio State University 1735 Neil Avenue Columbus, Ohio 43210/USA

W.P. JACOBS Department of Biology Princeton University Princeton, New Jersey 08544/USA

H. KALDEWEY

Fachrichtung Botanik Universitat des Saarlandes 6600 Saarbrucken/FRG

A.C. LEOPOLD Boyce Thompson Institute Ithaca, New York 14853/USA

A.G. MATTHYSSE Department of Biology University of North Carolina Chapel Hill, North Carolina 27514/USA

A.W. NAYLOR Department of Botany Duke University Durham, North Carolina 27706/USA

L.D. NOODEN Botany Department University of Michigan Ann Arbor, Michigan 48109-1048/USA

J .A. ROMBERGER

Light and Plant Growth Laboratory Institute of Plant Physiology BARC-West, Bldg. 046 A Beltsville, Maryland 20705/USA

T.K. SCOTT Department of Biology The University of North Carolina Chapel Hill, North Carolina 27514/USA

T.J. WODZICKI Department of Forest Botany Warsaw Agricultural University 26/30 Rakowiecka Street 02-528 Warsaw/Poland

S. ZAJ4CZKOWSKI Department of Forest Botany Warsaw Agricultural University 26/30 Rakowiecka Street 02-528 Warsaw-Poland