Molecular Genetics of. Plant Development STEPHEN H. HOWELL CAMBRIDGE UNIVERSITY PRESS

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1 Molecular Genetics of Plant Development STEPHEN H. HOWELL CAMBRIDGE UNIVERSITY PRESS

2 Contents Preface A Word on Genetic Nomenclature page xiii xvii 1 Approaches to the Study of Plant Development 1 Pattern Formation in Development 4 Important Differences between Plant and Animal Development 9 Germ Line Development 9 Role of the Gametophyte 10 Postembryonic Development 11 Cell Movement and the Planes of Cell Division 11 Regeneration and Totipotency 13 Variety of Plant Organs and Cell Types 15 Intercellular Communication 16 Model Plant Systems 18 Cloning Genes Relevant to Plant Development 20 The "Virtual Plant" 24 References 24 2 Cell Lineages and Positional Information 27 Predictability of Cell Fates 27 Marking Cells within a Lineage 28 Founder Cells and Developmental Compartments 28 Determining the Number of Founder Cells by Cell Lineage Analysis 30 Cell Fate Mapping in Early Embryogenesis 35 Experimental Disruption of Cell Lineages 38 Positional Information in Animal Development 38 Positional Information in Plant Development 40 Cell Layers and Chimeras 44 Transmission of Information between Cell Layers 48 Knotted Gene 48 vn

3 Floricaula Gene 50 References 53 3 Embryogenesis 55 Early Events in Embryogenesis 5 5 Complexity of Gene Expression in the Embryo 59 Genetics of Embryogenesis 60 Embryo-Lethal Mutants 61 Arabidopsis 61 Maize 64 Pattern Mutants 64 Polarity or Apical-Basal Axis Mutants 66 Short Integument Mutants 66 Gnom/emb30 Mutants 67 Segment Deletion Mutants 69 Monopteros Mutants 69 Gurke Mutants 71 Radial Axis Mutants 71 Knolle Mutants 71 Summary about Pattern Formation in Embryogenesis 72 Cell Fate Maps in Embryo Development 73 In vitro Fertilization and Development of Plant Embryos 75 Somatic Embryogenesis 76 References, 80 4 Seedling Development 83 Photomorphogenesis 84 Light-Insensitive Photomorphogenic Mutants - hy Mutants 85 Constitutive Light Response Mutants - det and cop Mutants 88 Fusca Mutants 92 Downstream Events in Photomorphogenesis 93 Brassinosteroids and the Control of Photomorphogenesis 94 Ethylene Mutants and the Triple Response 96 References Shoot Development 103 SAM Organization - Layers and Zones 104 Vlll

4 SAM Mutants 108 Mutants That Fail to Maintain Proliferative Cells in the Meristem 108 Mutants That Overproduce Proliferative Cells in the Meristem 114 Cell Determination in the SAM 117 SAM Molecular Biology 119 Phyllotaxy 121 Elongation of Internodes and Gibberellic Acid 127 Phase Changes in Vegetative Growth 129 References 133 Leaf D evelopment 136 Leaf Determination 137 Origins of Leaf Primordia in Dicots 139 Dicot Leaf Development 143 Genetics of Leaf Development 146 Dicots 146 Monocots 150 Compound Leaves 154 Trichome Development and Patterning 157 Stomatal Patterning and Spacing 162 References 166 Transition to Flowering 169 Transition to Flowering in Pea 170 Transition from Vegetative to Inflorescence Development in Arabidopsis 173 Transition from Inflorescence to Floral Development in Arabidopsis 178 Control of Flowering through the Action of Transgenes 181 Transition from Inflorescence to Floral Development in Antirrhinum 184 Floral Transition in Noninducible Systems 187 References 190 Flower Development 192 Founding Cell Populations for Flowers 194 Molecular Genetics and the ABC Model 195 Identification of Floral Homeotic Genes 200 IX

5 Cadasteral Functions of Homeotic Genes 201 Positive Regulation of Homeotic Gene Function 204 Molecular Functions of MADS Box Genes 207 Genes That Mediate the Interactions between Floral Meristem and Floral Organ Identity Genes 210 Control of Whorl Formation and Number of Organs per Whorl 213 Sex Determination - Monecious and Dioecious Plants 216 References Development of Floral Reproductive Organs and Gametophytes 222 Anther Development and Pollen Formation 224 Genetics of Anther Development and Pollen Formation 228 Nuclear Male Sterility 229 Cytoplasmic Male Sterility 231 Pistil, Ovule, and Embryo Sac Development 232 Mutants That Affect Pistil, Ovule, and Embryo Sac Development 238 References Pollination and Apomixis 244 Pollen Germination and Pollen Tube Growth 246 Pollen Tube Guidance 248 Self-Incompatibility 252 Sporophytic Self-Incompatibility 254 Gametophytic Self-Incompatibility 256 Apomixis 258 References Seed and Fruit Development 263 Seed Development 263 Morphogenesis Stage 265 Development of the Endosperm 265 Maturation Stage 270 Desiccation Stage 274 Dormancy and the Control of Germination 275 VP1 Gene in Maize 275 ABI-3, LEC1, and FUS3 Genes in Arabidopsis 277 Fruit Development 280 Fruit Growth 280 Fruit Ripening 282 References 285

6 12 Root Development 289 The Root Apical Meristem and the Promeristem 290 Cell Fates and Cell Lineage Analysis 292 The Role of Positional Information in Root Development 295 The Origin of the Primary Root during Embryogenesis 297 Lateral Roots 300 Genetics of Root Development 303 Root Hairs 306 References Vascular Development 312 Cells of the Vasculature 312 Development of the Stem and Leaf Vasculature 316 Regeneration of Vascular Strands 320 Lineage of Vascular Cells in the Leaf 322 Differentiation of Xylem and Phloem and Programmed Cell Death 325 Gene Expression during Vascular Differentiation 327 Mutants That Affect Vascular Development 331 References 334 Index 337 XI

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