Apomixis in Plants. Authors. Sven E. Asker, Ph.D. Department of Genetics University of Lund Lund, Sweden

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1 Apomixis in Plants I (0 ') r,\ q f Authors Sven E. Asker, Ph.D. Department of Genetics University of Lund Lund, Sweden Lenn Jerling, Ph.D. Botany Department University of Stockholm Stockholm, Sweden CRC Press Boca Raton Ann Arbor London Tokyo

2 TABLE OF CONTENTS Chapter 1 Introduction 1 References 2 Chapter 2 Apomixis and Parthenogenesis A Historical Review 5 I. The Empirical Background 5 A. Early Observations 5 B. Mendel on Apomixis 7 C. Pioneer Work 8 D. Biosystematic and Biogeographic Observations 8 II. Early Theories for Apomixis 11 A. The Necrohormone Theory 11 B. The Hybridization Theory 11 ' C. Gustafsson's Monograph 13 III. Advances since Gustafsson's Monograph 14 A. Genetic Background of Apomixis 14 B. Advances in Embryology and Cytology 15 C. Techniques for Studies of Variation in Apomicts 16 D. Apomixis vs. Sexual Reproduction Progress in General Theory 16 IV. Aspects of Apomixis 17 A. Ecological Interest in Apomicts Increases 17 B. The Evolutionary Potential of Apomicts 17 C. Plant Breeding 19 References 19 Chapter 3 ; Parthenogenesis in Animals 25 I. Introduction 25 II. Classification of Animal Parthenogenesis 25 III. Artificial and Occasional (Tycho-) Parthenogenesis 26 IV. Parthenogenesis and Polyploidy in Animals 27 V. Cyclical Parthenogenesis 29 VI. Evolution of Parthenogenetic Animals 31 VII. Comparison between Apomixis and Parthenogenesis in Plants and Animals 31 References 33 Chapter 4 Parthenogenesis, Apospory, and Apogamy in Lower Plants 37 I. Introduction 37

3 Chapter 8 Variation and Evolution of Apomicts 121 I. The Evolutionary Potential of Apomicts : 121 A. The Theory of Lack of Variation 121 B. Accepting Genetic Variation in Apomicts Generation of Variability in Apomicts Evolutionary Prospects 123 II. Empirical Studies 124 A. Biometrics 124 B. Biochemical Studies of Variation: DNA, Allozymes, Secondary Compounds DNA Studies Allozyme Studies Secondary Compounds 129 III. Further Aspects of Evolutionary Potential 129 A. Intraindividual Selection 129 B. Phenotypic Plasticity 132 C. Recombination and DNA Repair 132 D. The Speciation Rate 132 E. Sex as a Cohesive Force 133 IV. Empirical Records 133 V. Conclusion 134 References 135 Chapter 9 Taxonomic Considerations 139 I. Taxonomy The Problem with Classification 139 A. Introduction 139 B. Species Concepts The Biological Species The Phenetic Species Species as Evolutionary or Ecological Groups Species as Phylogenetic Groups 147 II. Taxonomical Trends 150 III. Conclusion 157 References 158 Chapter 10 Some Theoretical Considerations of Sexual Reproduction and Genetic Variation 161 I. The Process of Living 161 A. Introduction 161 B. The Fitness Concept 165 C. Cost of Sexual Reproduction 166

4 II. Selected Hypotheses for the Origin and Maintenance of Sexual Reproduction 167 A. Theories within POL Recombination as a Result of the Need for DNA Repair Clones vs. Clades Arguments for Sexual Reproduction Based on Speciation Rates Sex as a Consequence of the Level of Organization 169 B. Mendelian and Neo-Darwinian Hypotheses for Sex The Vicar of Bray The Ratchet The Best Man The Tangled Bank The Red Queen 172 References 173 Chapter 11 Ecological Implications of Apomixis 177 I. Introduction 177 II. Some Demographical Considerations 177 III. Two General Interpretations of Apomixis 180 A. Spreading the Risk of Mortality 180 B. Apomixis in Sparse Populations Stochasticity Density-Dependent Effects 183 IV. Some Ecological Implications of Polyploidy 184 V. Competition 185 A. Inter- or Intraspecific Competition? i 186 B. Effects of Competition on the Individual 186 C. Competition and the Relative Production of Sexual and Apomictic Seeds 187 VI. Coexistence 188 A. Introduction 188 B. Equilibrium Theories Coexistence between Clones Coexistence between a Sexual and an Asexual Type Frequency Dependence 192 C. Nonequilibrium Theories Fugitives and Dominants Environmental Fluctuations and Buffering Slow Competitive Displacement 193 D. Continuous Recruitment of New Clones 194

5 VII. Herbivory and Apomictic Plants 194 VIII. Pathogens and Apomictic Plants 196 IX. Ecotype Differentiation in Apomictic Plants 198 A. The Process 198 B. The Raw Material Genetic Variation 199 C. Effects of Apomixis on Gene Flow 200 D. Effects of Apomixis on Genetic Drift 201 E. Apomixis and Selection 201 F. Apomixis, Phenotypic Plasticity, and Response to Selection 203 G. Some Aspects on the Relation between Apomixis and Natural Selection 204 X. Aspects of Life History Patterns in Apomicts 205 A. Introduction 205 B. Life Form, Size, Vegetative Propagation, and Mobility 206 C. Rate of Growth and Development 207 D. Reproduction, Sexual System, and Pollination 207 E. Flowering Time and Phenology 208 F. Fecundity, Seed Size, and Germination 209 G. Habitats of Apomicts 210 References 211 Chapter 12 Geographical Trends 221 I. Patterns of Distribution 221 A. Latitude 221 B. Distributional Patterns of Apomicts and Amphimicts within a Species 222 C. Levels of Altitude 228 D. Occupancy of Glaciated Areas 228 II. Historical Aspects Distribution and Age of Apomicts 230 A. General Remarks 230 B. Some Examples Crepis Rubus Taraxacum Hieracium Citrus 234 C. Classification of Apomictic Complexes 235 III. Conclusion 236 References 237

6 Chapter 13 Apomixis and Breeding 241 I. Introduction 241 II. Breeding Apomictic Crops 242 A. Background 242 B. Cultivated or Domesticated Species Which Normally Have Apomictic Reproduction 243 C. Fruit Trees and Berry Bushes with Apomixis Citrus Other Fruit Trees and Berry Bushes 244 D. Fodder Grasses with Apomixis Panicum maximum Other Tropical and Subtropical Forage Grasses Poa pratensis 247 E. "Rubber Plants" of Asteraceae (Compositae) 248 III. Use of Apomixis in Breeding Crops with Sexual or Vegetative Reproduction 249 A. The Approach: "Transfer" vs. "Synthesis" 249 B. Maize 251 C. Rice 254 D. Wheat 254 E. Barley 255 F. Sorghum 256 G. Pennisetum 257 H. Distant Hybridization in Fodder Grasses 258 I. Sugar Beet 258 J. Brassica and Raphanobrassica 259 K. Strawberry 260 L. Apple ; 260 M. Cherry 261 N. Pea, Soybean, Alfalfa, and other Fabaceae 261 O. Cotton 261 P. Potato 261 Q. Tobacco 262 R. Other Crop Plants 262 S. Some Final Words 263 References 263 Glossary 279 Index 285

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