Pollination Mechanisms, Reproduction and Plant Breeding

293 Pages·1977·8.49 MB·English

by  Professor Dr. Rafael Frankel

Monographs on
2
Theoretical and Applied Genetics
Edited by
R. Frankel (Coordinating Editor), Bet Dagan
G. A. E. Gall, Davis . M. Grossman, Urbana
H. F. Linskens, Nijmegen . D. de Zeeuw, Wageningen
R. Frankel . E. Galun
Pollination Mechanisms,
Reproduction
and Plant Breeding
With 77 Figures
Springer-Verlag
Berlin Heidelberg New York 1977
Professor Dr. RAFAEL FRANKEL, Agricultural Research Organization,
The Volcani Center, P. O. Box 6, Bet Dagan/Israel
Professor Dr. ESRA GALUN, Weizman Institute of Science, Rehovot/
Israel
ISBN-13 :978-3-642-81 061-9 e-ISBN-13:978-3-642-81059-6
DOl: 10.1007/978-3-642-81059-6
Library of Congress Cataloging in Publication Data. Frankel, Rafael, 1922. Pollination mecha·
nisms, reproduction, and plant breeding. (Monographs on theoretical and applied genetics;
2) Bibliography: p. Includes index. 1. Plant-breeding. 2. Fertilization of plants. 3. Plants-Repro
duction. r. Galun, E., 1927. joint author. II. Title. III. Series. SB123.F7 631.5'3.76-42450.
This work is subject to copyright. All rights are reserved, whether the whole or part of the material
is concerned, specifically those of translation, re-use of illustrations, broadcasting, reproduction
by photocopying machine or similar means, and storage in data banks. Under § 54 ofthe German
Copyright Law, where copies are made for other than private use, a fee is payable to the publisher,
the amount of the fee to be determined by agreement with the publisher.
© by Springer-Verlag Berlin Heidelberg 1977.
Softcover reprint of the hardcover 1st edition 1977
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 protectice laws and
regulations and therefore free for general use.
Preface " ... Nature has something more in
view than its own proper males should
fecundate each blossom."
ANDREW KNIGHT
Philosophical Transactions, 1799
Pollination mechanisms and reproduction have a decisive
bearing upon rational procedures in plant breeding and crop
production. This book intends to furnish' under one cover
an integrated botanical, genetical and breeding-methodologi
cal treatment of the reproductive biology of spermatophytes
mainly angiosperms; it is based on an advanced topical course
in plant breeding taught at the Hebrew University of Jerusa
lem.
We have tried to present a coverage which is concise,
but as comprehensive as possible, of the pollination
mechanism and modes of reproduction of higher plants, and
to illustrate topics, whenever practicable, by examples from
cultivated plants. Nevertheless, some relevant publications
may have escaped our attention or may not be mentioned
because of various limitations.
The book is organized into three parts. The first part
starts with an evaluation of the significance of the different
pollination mechanisms for plant breeding and crop produc
tion, describes modes of reproduction in higher plants and
discusses ecology and dynamics of pollination. The second
part is devoted to crops propagated by self pollination and
describes specific breeding procedures for such crops. The
third part details sexual reproduction in higher plants and
handles three mechanisms involved in the prevention of self
pollination and their utilization in plant breeding: sex expres
sion, incompatibility, and male sterility.
We hope this book will prove useful to biology and agricul
ture students at the graduate level, to plant breeders and
other people interested in acquiring a broader knowledge
of the reproductive biology of higher plants, and serve for
intra-and interdisciplinary communication between botanists,
geneticists, plant breeders and agriculturists.
We owe thanks to Mrs. V. ASSCHER for her careful
typing of the manuscript, to Mrs. V. PRIEL for English editing,
and to Mrs. N. GESTETNER for preparing the line drawings.
One of us (E. G.) is grateful to the Israel Academy of Sciences
VI Preface
and Humanities and to the Royal Society for a grant which
enabled him to perform a literature study at the Royal Botanic
Gardens, Kew. Thanks are also due to many colleagues and
students for valuable discussions and suggestions and to
authors and publishers for permission to use or reproduce
their illustrations.
Bet Dagan/Rehovot, R. FRANKEL . E. GALUN
January 1977
Contents
Chapter 1. Introduction 1
1.1 Implications of Pollination Mechanisms in Plant
Breeding and Crop Production . . . . . . 1
1.1.1 Pollination Mechanisms and Breeding of New Culti-
vars .................... . 1
1.1.2 Pollination Mechanisms and Cultivar Maintenance. 2
1.1.3 Pollination Mechanisms and Agricultural Yield 3
1.2 Reproduction in Higher Plants . . . . . 3
1.2.1 Historical and General Background . . . 3
1.2.1.1 Morphological-Structural Flower Biology . 4
1.2.1.2 Functional Flower Biology . . . . . . . 4
1.2.1.3 Correlation between Structure and Function. 5
1.2.2 Modes of Reproduction . . . . . . . . . 5
1.2.2.1 Sexual Reproduction. . . . . . . . . . . 6
1.2.2.1.1 Spatial Separation of Sex Organs as an Outbreeding
Device ................ . 6
1.2.2.1.2 Basic Concepts of Structural Differentiation . 6
1.2.2.1.3 Models of Floral Differentiation. . . . . . 7
1.2.2.1.4 Definitions of Sex Types in Flowering Plants 10
1.2.2.1.5 Temporal Separation of Sex Organs as an Outbreeding
Device ................ . 13
1.2.2.2 Asexual Forms of Reproduction. . . . . . 14
1.2.2.2.1 Asexual Propagules outside the Floral Region 14
1.2.2.2.2 Asexual Propagules within the Floral Region 15
1.2.2.3 Distribution of Modes of Reproduction among Culti-
vated Plants. . . . . . . . . . . . 16
1.3 Ecology and Dynamics of Pollination. 29
1.3.1 Specificity of Flowers and Pollen. 29
1.3.2 Pollen-Dispersal Agents 31
1.3.2.1 Biotic Vectors. . . . . . . . . 31
1.3.2.2 Abiotic Vectors . . . . . . . . 34
1.3.3 Timing and Climatic Factors in Pollination Dynamics 36
1.3.4 Location and Mass Effects in Pollination Ecology . 41
1.3.5 Competition Effects in Pollination Dynamics . . . 43
1.3.6 Determination of the Natural Cross-Pollination Rate
(NCP) .......... . 43
1.3.6.1 Progeny Testing of Dominants. 45
1.3.6.2 Progeny Testing of Recessives . 46
VIII Contents
1.3.6.3 Progeny Testing of Heterozygotes . . . . . . 46
1.3.6.4 Progeny Testing of Recessives and Heterozygotes. 47
1.3.7 Artificial Control of Outcrossing . 48
Chapter 2. Autogamy ........ 51
2.1 Evolutionary Aspects of Autogamy. 51
2.1.1 Strategies for Adjustment of Recombination . 51
2.1.2 Origin of Mating Systems in Higher Plants 52
2.1.3 Variation in Autogamous Populations 53
2.2 Mechanism of Autogamy . 56
2.2.1 Cleistogamy........... 56
2.2.2 Chasmogamic Selfing . . . . . . . 58
2.3 Management of Pollination in Autogamous Crops 62
2.3.1 Emasculation............. 62
2.3.1.1 Mechanical Removal of Microsporophylls . . . 62
2.3.1.2 Male Gametocide . . . . . . . . . . . . . 66
2.3.1.3 Circumvention of Emasculation Requirements. 69
2.3.2 Controlled Pollination . . . . 70
2.3.2.1 Isolation.......... 70
2.3.2.2 Pollen Collection and Storage . 70
2.3.2.3 Pollen Transfer Methods . . . 73
2.3.2.3.1 Utilization of Natural Pollen Vectors. 73
2.3.2.3.2 Forced Pollination. 76
Chapter 3. Allogamy 79
3.1 Sexual Reproduction-Structures and Functions. 79
3.1.1 The Anther and the Male Gametophyte 79
3.1.1.1 Differentiation of the Anther . . . . . . 79
3.1.1.2 The Tapetum. . . . . . . . . . . . 80
3.1.1.3 Development of the Sporogeneous Tissue 81
3.1.1.4 Microsporogenesis.......... 82
3.1.1.5 From Microspore to Pollen Grain . . . 83
3.1.2 Androgenesis: Production of Haploid Plants by
Anther and Pollen Culture . . . . . . . . . . . 85
3.1.2.1 Haploid Plants-Occurrence, Induction, and Identi
fication . . . . . . . . . . . . . . . . . . . . 85
3.1.2.2 The Production of Haploid Callus and Embryoids by
Anther and Pollen Culture . . . . 86
3.1.2.3 Pathways of Pollen Embryogenesis. 89
3.1.2.4 Factors Affecting Androgenesis 92
3.1.2.4.1 Culture Conditions 92
3.1.2.4.2 Donor Plants . . . . . 92
3.1.2.4.3 Pollen Age . . . . . . 92
3.1.2.4.4 Anther Stage and Ploidy 93
3.1.2.5 Application of Androgenesis for Breeding and Genetic
Studies. . . . . . . . . . . . . . . . . . . . 93
Contents IX
3.1.3 The Pistil and the Female Gametophyte. 96
3.1.3.1 The Pistil. . . . . . . . 97
3.1.3.2 The Female Gametophyte. . . . . 98
3.1.4 Fertilization . . . . . . . . . . 100
3.1.4.1 Contact between Pollen and Stigma 100
3.1.4.2 Pollen Germination . . . . . . . 101
3.1.4.3 Pollen Tube Discharge and Double Fertilization 102
3.1.5 Sexual Reproduction in Conifers. . . . . 103
3.2 Control and Modification of Sex. . . . . 104
3.2.1 The Genetic Control of Sex Determination 104
3.2.1.1 General Considerations Concerning the Genetic
Control of Dioecism. . . . . . . . . . . . . . 105
3.2.1.2 Artificial Dioecism in Zea mays . . . . . . . .. 107
3.2.1.3 Artificial Dioecism Caused by Suppressive Genes
Linkage between Genes as a Prerequisite for Sex
Dimorphism . . . . . . . . . . . . . . . . . 109
3.2.1.4 Chromosomal Control of Sex Determination . . . 110
3.2.1.5 Main Methods for the Study of the Genetics and
Cytology of Sex Determination . . . . . . . . . 112
3.2.1.6 Genetic Regulation of Sex: Representative Examples 113
3.2.1.6.1 The Active Y Chromosome System. . . . . . . . 113
3.2.1.6.2 The X-Autosomal Balance System. . . . . . . . 118
3.2.1.7 Genetics of Sex Determination in Some Economic
Crops .......... . 119
3.2.2 Modifications of Sex Expression. . . . 120
3.2.2.1 Introduction . . . . . . . . . . . . 120
3.2.2.2 Mineral Nutrition and Edaphic Factors. 127
3.2.2.3 Light . . . . . 128
3.2.2.4 Temperature . . . . . . . . . 132
3.2.2.5 Chemical Agents . . . . . . . 134
3.2.2.5.1 Auxins and Related Compounds. 135
3.2.2.5.2 Gibberellins. . . . . . . . . . 137
3.2.2.5.3 Kinins ........... . 139
3.2.3 Sex Expression in Some Economic Crops and its
Application to Breeding and Crop Improvement 141
3.2.3.1 The Cucumber and Other Cucurbit Crops. . . . . 141
3.2.3.1.1 Patterns of Sex Expression . . . . . . . . . . . 141
3.2.3.1.2 Effects of Day Length and Temperature on Flowering
and Sex Expression . . . . . . . . . . . . . . 145
3.2.3.1.3 Effects of Growth Regulators . . . . . . . . . . 147
3.2.3.1.4 Inheritance of Sex Expression and Breeding Proce-
dures . . . . . . . . . . . . . . . 151
3.2.3.1.5 Scheme of Sex Expression in Cucumber. 157
3.2.3.2 Hemp (Cannabis sativa) . 157
3.2.3.3 Maize (Zea mays). . . . . . . . . . 159
3.2.3.3.1 Reproductive Morphology . . . . . . 160
3.2.3.3.2 Environmentally Induced Sex Modification 160
x Contents
3.2.3.3.3 Sex Modification by Chemical Agents 161
3.2.3.3.4 Inheritance of Sex Expression 163
3.3 Incompatibility . . . . . . . 163
3.3.1 Genetics ofIncompatibility . . 165
3.3.1.1 Gametophytic Incompatibility. 165
3.3.1.1.1 One Multiallelic S Locus 165
3.3.1.1.2 Two Multia:llelic S Loci. . . 169
3.3.1.1.3 Three or More S Loci . . . 171
3.3.1.2 Sporophytic Incompatibility 172
3.3.1.2.1 Heteromorphic Incompatibility 173
3.3.1.2.2 Homomorphic Incompatibility 176
3.3.2 Pollen-Pistil Interaction. . . . 179
3.3.2.1 Pollen Cytology and Pollen-Stigma Interaction. 179
3.3.2.2 Pollen Tube-Style Interaction . . . . . . .. 183
3.3.2.3 Pollen Tube-Ovule Interaction. . . . . . .. 184
3.3.3 Incompatibility, Crop Production, and Breeding 184
3.3.3.1 Transfer of Incompatibility into Cultivars . 185
3.3.3.2 Permanent Elimination of Incompatibility. 186
3.3.3.3 Surmounting the Incompatibility Barrier 188
3.3.3.3.1 Treatment of Pollen 188
3.3.3.3.2 Bud Pollination . . 188
3.3.3.3.3 Delayed Pollination 188
3.3.3.3.4 Heat Treatment . . 188
3.3.3.3.5 Surgical Techniques 189
3.3.3.3.6 Double Fertilization 189
3.3.3.3.7 Other Methods . . 189
3.3.3.4 Hybrid Seed Production 190
3.3.3.4.1 Characteristics of the Gametophytic and Sporophytic
Incompatibility Systems which are Relat~d to Hybrid
Seed Production ................ 190
3.3.3.4.2 Hybrid Seed Production in Brassica and Raphanus . 191
3.3.3.4.3 Problems in Breeding Aimed at Hybrid Seed Produc-
tion . . . . . . . . . . . . . . . . . . . . . 195
3.4 Male Sterility. . . . . . . . . . . . . . . . . 196
3.4.1 Gynodioecy: Male Sterility in Natural and Artificial
Populations. . . . . . . . . . . . . . . . . . 196
3.4.1.1 Male Sterility as an Outbreeding Mechanism in
Gynodioecious Species . . . . . . . . . . . . . 196
3.4.1.2 Nuclear Male Sterility Genes in Cultivars . . . . . 199
3.4.1.3 Male-Sterility-Inducing Plasmatypes in Cultivated
Plants . . . . . . . . . . . . . . . . . 201
3.4.1.4 Utilization of Gynodioecy in Plant Breeding.. 203
3.4.2 Inheritance of Male Sterility. . . . . . . .. 204
3.4.2.1 Genic Male Sterility (Mendelian Male Sterility) 204
3.4.2.2 Cytoplasmic and Gene-Cytoplasmic Male Sterility 206
3.4.3 Structural, Developmental, and Biochemical Charac
terization of Male Sterility . . . . . . . . . . . 209
Contents XI
3.4.3.1 Developmental Modifications Leading to Breakdown
in Microgametogenesis. . . . . . . . . . . . . 209
3.4.3.1.1 Modifications in the Structural Differentiation of the
Stamen. . . . . . . . . . . . . 210
3.4.3.1.2 Faulty Differentiation of the Anther 212
3.4.3.1.3 Breakdown in Microsporogenesis . 214
3.4.3.1.4 Abortion of the Microgametophyte. 216
3.4.3.1.5 Functional Male Sterility. . . . . 219
3.4.3.2 Biochemistry of Male Sterility. . . 219
3.4.3.3 Male Sterility Genes and Their Action 221
3.4.3.3.1 Site of Male Sterility Factors . . . . 221
3.4.3.3.2 Pleiotropic Effects. . . . . . . . . 224
3.4.4 . Utilization of Male Sterility in Plant Breeding 225
3.4.4.1 Comparison of Hybrid Production Using Genic,
Cytoplasmic and Gene-Cytoplasmic Male Sterility 226
3.4.4.1.1 Genic Male Sterility . . . . . . 226
3.4.4.1.2 Cytoplasmic Male Sterility . . . . 227
3.4.4.1.3 Gene-Cytoplasmic Male Sterility. . 227
3.4.4.2 Application of Genic Male Sterility. 228
3.4.4.2.1 Genetic and Field Management Programs to Provide
a Homogeneous Stand of the Genic Male Sterile Seed
Parent. . . . . . . . . . . . . . . 228
3.4.4.2.2 Pollination Control . . . . . . . . . 230
3.4.4.3 Use of Gene-Cytoplasmic Male Sterility. 233
References . . 235
Subject Index 269