Table Of ContentEncyclopedia 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 NewY ork 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).
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© 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 scien
tific 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 labori
ous 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 ANDRE PIRSON
Spring 1984
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 knowl
edge 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 wide
reaching 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 compre
hensive 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 regula
tion, 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 organi
zation 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 ....... . . . . . . . . . . . . . . . . 4
1.1.1 The Concept of the Plant Hormone as a Chemical Messenger 4
1.1.2 Types of Hormones ............. . 5
1.2 Hormone Messenger Systems . . . . . . . . . . . . . 6
1.2.1 Common Difficulties with Single Hormonal Controls 6
1.2.2 Is a Hormone Always a Hormone? . . . . . . 7
1.2.3 Further Complications in the Hormone Concept 8
1.2.4 Regulatory Sites in Hormonal Controls 9
1.2.5 Patterns in Multiple Hormonal Control 10
1.2.6 Complications Arising from Compartmentation 11
1.3 Analysis of Hormonal Controls . . . . . 11
1.3.1 Two Major Approaches from the Past 12
1.3.2 Analyses of Hormone Systems 13
1.4 Conclusions ............ . 15
1.4.1 The Complexity of Hormonal Controls 15
1.4.2 Nonhormonal Correlative Controls . . 15
1.4.3 Evolutionary Plasticity of Hormonal Controls 16
1.4.4 The Future 16
References . . . . . . . . . . . . . . . . . . . . 17
2 Functions of Hormones at the Cellular Level of Organization
M.L. EVANS (With 1 Figure)
2.1 Introduction .. . . . . . . . . . . . . . . . 23
2.2 Promotive and Inhibitory Classes of Plant Hormones 23
2.3 Modes of Hormone Interaction . . . . . . . . . 24
2.3.1 Interaction and Independence in Hormone Action 24
2.3.2 Synergism and Antagonism in Hormone Action 25
2.3.2.1 Synergism ............ . 25
2.3.2.2 Antagonism ........... . 25
2.3.3 Hormone Effects on Endogenous Hormone Levels 27
2.3.3.1 Hormona: Modification of Endogenous Auxin 27
2.3.3.2 Hormonal Modification of Endogenous Gibberellin 28
2.3.3.3 Hormonal Modification of Endogenous Ethylene 29
2.3.3.4 Hormonal Modification of Endogenous Abscisic Acid 29
2.3.3.5 Hormonal Modification of Endogenous Cytokinin . . 30
2.3.3.6 Summary .................. . 30
2.3.4 Influence of Environmental Parameters on Endogenous Hormones 31
2.3.4.1 Light 31
2.3.4.2 Water Stress 33
XII Contents
2.3.4.3 Temperature . . . . . 34
2.3.4.4 Mechanical Stimulation 34
2.3.4.5 Summary 34
2.4 Hormonal Regulation . . . . . 35
2.4.1 The Cell Cycle . . . . . . 35
2.4.1.1 Effect of Cytokinins 36
2.4.1.2 Effect of Auxins . . 39
2.4.1.3 Effects of Other Growth Regulators 40
2.4.2 Endomitosis 42
2.4.3 Meiosis . . . . . . . . . . . . . . 43
2.4.4 Summary . . . . . . . . . . . . . 44
2.5 Hormone Effects on the Plane of Cell Division 44
2.6 Relationship of Cell Multiplication to Cell Expansion 46
2.7 Hormonal Regulation of Cell Expansion 48
2.7.1 Stem Cells ....... . 48
2.7.1.1 Effects of Auxin . . . . . 48
2.7.1.2 Effects of Gibberellin 49
2.7.1.3 Effects of Cytokinin, Ethylene, and Abscisic Acid 49
2.7.2 Root Cells ....... . 50
2.7.2.1 Effects of Auxin . . . 51
2.7.2.2 Effects of Gibberellin 52
2.7.2.3 Effects of Ethylene 53
2.7.2.4 Effects of Abscisic Acid 54
2.7.2.5 Effects of Cytokinin 54
2.7.3 Leaf Cells . . . . . . . . 55
2.7.3.1 Effects of Auxin . . 55
2.7.3.2 Effects of Gibberellin 56
2.7.3.3 Effects of Ethylene 56
2.7.3.4 Effects of Abscisic Acid 56
2.7.3.5 Effects of Cytokinin 56
2.7.4 Other Cell Types . . . 57
2.7.4.1 Pollen Tubes 57
2.7.4.2 Epidermal Hairs 57
2.7.5 Lower Plants 57
2.7.5.1 Algae 57
2.7.5.2 Fungi 59
2.7.6 Summary 61
References . . . . . . 62
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
