Seed development, dormancy and germination /
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作 者：edited by Kent J. Bradford and Hiroyuki Nonogaki.
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ISBN：9781405139830
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简介
Summary:
Publisher Summary 1
This overview of seed biology covers the development and regulatory process from the developing seed to dormancy to germination and seedling growth, keeping in mind the complex environmental, physiological, molecular and genetic interactions that occur throughout the cycle. The 12 papers here cover the genetic control of seed development and seed mass, seed coat development and dormancy, definitions and hypotheses about seed dormancy, seed dormancy (modeling, genetic aspects, lipid metabolism, abscisic acid and hormonal cross-talk) nitrous oxide in seed dormancy and germination, regulation of ABA and GA levels, DE-repression of seed germination by GA signaling, mechanisms and genes involved in germination, and the role of sugar and abscisic acid regulation in germination and the transition to seedling growth. Annotation 漏2007 Book News, Inc., Portland, OR (booknews.com)
Publisher Summary 2
The formation, dispersal and germination of seeds are crucial stages in the life cycles of gymnosperm and angiosperm plants. The unique properties of seeds, particularly their tolerance to desiccation, their mobility, and their ability to schedule their germination to coincide with times when environmental conditions are favorable to their survival as seedlings, have no doubt contributed significantly to the success of seed-bearing plants. Humans are also dependent upon seeds, which constitute the majority of the world's staple foods (e.g., cereals and legumes). Seeds are an excellent system for studying fundamental developmental processes in plant biology, as they develop from a single fertilized zygote into an embryo and endosperm, in association with the surrounding maternal tissues. As genetic and molecular approaches have become increasingly powerful tools for biological research, seeds have become an attractive system in which to study a wide array of metabolic processes and regulatory systems.
Seed Development, Dormancy and Germinationprovides a comprehensive overview of seed biology from the point of view of the developmental and regulatory processes that are involved in the transition from a developing seed through dormancy and into germination and seedling growth. It examines the complexity of the environmental, physiological, molecular and genetic interactions that occur through the life cycle of seeds, along with the concepts and approaches used to analyze seed dormancy and germination behavior. It also identifies the current challenges and remaining questions for future research. The book is directed at plant developmental biologists, geneticists, plant breeders, seed biologists and graduate students.
目录
Table Of Contents:
List of Contributors xiii
Preface xv
1 Genetic control of seed development and seed mass 1
MASA-AKI OHTO, SANDRA L. STONE AND JOHN J. HARADA
1.1 Introduction 1
1.2 Overview of seed development in angiosperms 1
1.3 Genetic control of embryo development 3
1.3.1 Central regulators of embryogenesis 3
1.3.2 Genes involved in the morphogenesis phase of embryo development 4
1.3.3 Regulators of the maturation phase of embryo development 5
1.4 Genetic control of endosperm development 6
1.4.1 Genes required for cereal endosperm development 7
1.4.2 Genes that repress autonomous endosperm development 7
1.5 Genetic aspects of testa development 8
1.5.1 Genetic regulation of flavonoid biosynthesis and accumulation 9
1.5.2 Regulators of mucilage biosynthesis and accumulation 9
1.6 Control of seed mass 10
1.6.1 Genetic factors affecting seed mass 10
1.6.2 Testa development and seed mass 11
1.6.3 Endosperm development and seed mass 11
1.6.4 Sugar transport and metabolism during seed development 13
1.6.5 Metabolic control of seed development and size 15
1.7 Perspective 17
References 17
2 Seed coat development and dormancy 25
ISABELLE DEBEAUJON, LOIC LEPINIEC, LUCILLE POURCEL AND JEAN-MARC ROUTABOUL
2.1 Introduction 25
2.2 Development and anatomy of the seed coat 25
2.2.1 The seed envelopes 25
2.2.2 The Arabidopsis testa 27
2.3 Role of the seed coat in seed dormancy and germination 29
2.3.1 Constraints imposed by the seed coat 29
2.3.2 Flavonoids in Arabidopsis seeds 31
2.3.2.1 Main flavonoid end-products present in seeds 31
2.3.2.2 Molecular genetics of flavonoid metabolism 32
2.3.2.3 Effects of flavonoids on seed dormancy and germination 35
2.3.3 Flavonoids in seed dormancy and germination of various species 38
2.3.3.1 Solanaceae 38
2.3.3.2 Water permeability of testae in Leguminosae and other species 39
2.3.3.3 Flavonoids and other phenolics as direct and indirect germination inhibitors 39
2.3.3.4 Pre-harvest sprouting (PHS) in cereals 40
2.3.3.5 Heteromorphism and physiological heterogeneity among seeds 40
2.3.3.6 Interactions with endosperm 41
2.4 Link between seed coat-imposed dormancy and longevity 41
2.5 Concluding remarks 42
References 43
3 Definitions and hypotheses of seed dormancy 50
HENK W.M. HILHORST
3.1 Introduction 50
3.2 Classifications of dormancy 50
3.2.1 Endogenous dormancy 52
3.2.2 Exogenous dormancy 53
3.3 Definitions of dormancy 54
3.4 Primary dormancy 57
3.4.1 Induction of primary dormancy 57
3.4.1.1 Role of ABA in dormancy induction 57
3.4.1.2 Developmental programs and dormancy induction 58
3.4.2 Release of primary dormancy 59
3.4.2.1 After-ripening 59
3.4.2.2 Regulation of dormancy in imbibed seeds 60
3.5 Secondary dormancy 63
3.6 Signaling in dormancy 64
3.6.1 Stress signaling 64
3.6.2 Signaling networks 65
3.6.3 Environmental signals 65
3.7 Challenges for the future 67
References 67
4 Modeling of seed dormancy 72
PHIL S. ALLEN, ROBERTO L. BENECH-ARNOLD, DIEGO BATLLA AND KENT J. BRADFORD
4.1 Introduction 72
4.2 Types and phenology of seed dormancy 73
4.3 Environmental control of dormancy 76
4.3.1 Factors affecting dormancy levels of seed populations 76
4.3.1.1 Temperature 76
4.3.1.2 After-ripening 78
4.3.1.3 Stratification 79
4.3.2 Factors that stimulate germination 80
4.3.2.1 Fluctuating temperature 80
4.3.2.2 Light 81
4.3.2.3 Nitrate 82
4.3.3 Conceptual scheme of dormancy and its relationship to modeling 82
4.4 Approaches to modeling seed dormancy 82
4.4.1 Temperature response models and thermal time 83
4.4.2 Water potential responses and hydrotime models 87
4.4.3 Interactions of temperature and water potential 89
4.4.4 Modeling responses to other factors affecting dormancy and germination 90
4.5 Examples of seed dormancy models 91
4.5.1 Solanum tuberosum 91
4.5.2 Bromus tectorum 92
4.5.3 Polygonum aviculare 96
4.5.3.1 Modeling seed germination responses to temperature 97
4.5.3.2 Modeling seed responses to germination-stimulating factors 100
4.6 Population-based threshold models of seed dormancy 100
4.7 Conclusions and future directions 105
References 106
5 Genetic aspects of seed dormancy 113
LE脫NIE BENTSINK, WIM SOPPE AND MAARTEN KOORNNEEF
5.1 Introduction 113
5.2 Mutant approaches in Arabidopsis 114
5.3 Mutant approaches in other species 118
5.4 Genetic analyses of natural variation 119
5.4.1 Genetic analysis of natural variation in Arabidopsis 120
5.4.2 Natural variation for dormancy in grasses 120
5.5 What do the genetics teach us about dormancy and germination? 125
References 127
6 Lipid metabolism in seed dormancy 133
STEVEN PENFIELD, HELEN PINFIELD-WELLS AND IAN A. GRAHAM
6.1 Introduction 133
6.2 Metabolic pathways for TAG breakdown and conversion to sucrose 135
6.2.1 TAG hydrolysis and activation 135
6.2.2 Import of fatty acids into the peroxisome 135
6.2.3 Activation of fatty acids to acyl-CoA thioesters for β-Oxidation 136
6.2.4 β-Oxidation 137
6.2.4.1 Acyl-CoA oxidases 137
6.2.4.2 Multifunctional protein 137
6.2.4.3 3-L-Ketoacyl-CoA thiolase 138
6.2.4.4 Peroxisomal citrate synthase 138
6.2.5 Glyoxylate cycle and gluconeogenesis 138
6.2.5.1 Isocitrate lyase 139
6.2.5.2 Malate synthase 139
6.2.5.3 Phosphoenolpyruvate carboxylase 139
6.3 Lipid metabolism and seed dormancy 140
6.3.1 Importance of the ABC transporter for the transition from dormancy to germination 140
6.3.2 Defects in β-oxidation enzymes, but not in LACS, affect seed dormancy 142
6.3.3 Storage lipid mobilization (glyoxylate cycle and gluconeogenesis) is not required for seed dormancy release 144
6.4 Mechanisms for the involvement of β-oxidation in dormancy release 145
6.4.1 β-Oxidation does not fuel seed germination 145
6.4.2 β-Oxidation and hormonal signaling 145
6.4.3 Possible biosynthetic roles for β-oxidation in regulating germination 147
6.4.4 β-Oxidation, reactive oxygen species, and redox control 148
6.5 Conclusions 149
References 149
7 Nitric oxide in seed dormancy and germination 153
PAUL C. BETHKE, IGOR G.L. LIBOUREL AND RUSSELL L. JONES
7.1 Nitric oxide in plant growth and development 153
7.2 Challenges in NO chemistry and biology 153
7.3 Tools used in NO research 154
7.4 Roles of NO and other N-containing compounds in seed dormancy and germination 157
7.4.1 Nitrate, nitrite, and ammonium 158
7.4.2 Cyanide and azide 159
7.4.3 NO donors and germination 160
7.4.4 NO scavengers and germination 162
7.5 Biochemical and molecular basis of NO action in seeds 162
7.5.1 Synthesis of NO by plants 162
7.5.2 NO binding to metal-containing proteins 164
7.5.3 NO as an antioxidant 166
7.6 Interactions between NO and phytochrome or ABA 167
7.7 Ecological significance of NO 168
7.7.1 Nitrogen and vegetation gap sensing 168
7.7.2 Smoke and NO 169
7.8 Unresolved questions and concluding remarks 169
References 171
8 A merging of paths: abscisic acid and hormonal cross-talk in the control of seed dormancy maintenance and alleviation 176
J. ALLAN FEURTADO AND ALLISON R. KERMODE
8.1 Introduction 176
8.2 Abscisic acid 177
8.2.1 ABA in seed maturation and the induction of primary dormancy 177
8.2.2 Transcription factors and combinatorial control of seed development and maturation 180
8.2.3 ABA in dormancy maintenance and termination 182
8.2.3.1 ABA synthesis and homeostasis during dormancy maintenance and termination 182
8.2.3.2 ABA signaling factors and the control of dormancy maintenance and termination 186
8.3 Gibberellin 190
8.3.1 GA is antagonistic to ABA during seed development 190
8.3.2 GA promotes the transition to germination 191
8.4 Light interactions 195
8.4.1 GA synthesis and signaling are promoted by light through the action of phytochrome 195
8.4.2 ABA-associated signaling processes are opposed by light signaling 196
8.5 Ethylene 197
8.5.1 Ethylene counteracts ABA during seed development 197
8.5.2 Ethylene promotes the transition from dormancy to germination 198
8.6 Auxin and cytokinin 201
8.6.1 Auxin and cytokinin establish the embryo body plan during seed development 201
8.6.2 Auxin and cytokinins have not been intimately linked to dormancy maintenance or termination 202
8.7 Brassinosteroids 203
8.8 G-protein signaling reveals integration of GA, BR, ABA, and sugar responses 205
8.9 Profiling of hormone metabolic pathways in Arabidopsis mutants reveals cross-talk 206
8.10 Summary and future directions 208
References 211
9 Regulation of ABA and GA levels during seed development and germination in Arabidopsis 224
SHINJIRO YAMAGUCHI, YUJI KAMIYA AND EIJI NAMBARA
9.1 Introduction 224
9.2 Biosynthetic and deactivation pathways of ABA and GA 225
9.2.1 ABA biosynthesis 225
9.2.2 ABA deactivation 229
9.2.3 ABA-deficient mutants and seed germination 230
9.2.4 GA biosynthesis 230
9.2.5 GA deactivation 231
9.2.6 GA-deficient mutants and seed germination 231
9.3 Inhibitors of ABA and GA metabolism: efficacy and side effects of drugs 232
9.3.1 Drugs to reduce endogenous ABA levels 232
9.3.2 Drugs to increase endogenous ABA levels 234
9.3.3 Drugs to reduce GA levels 235
9.3.4 Side effects of drugs 235
9.4 Regulation of ABA and GA levels in Arabidopsis seeds 236
9.4.1 Regulation of ABA and GA levels during seed development 236
9.4.1.1 Roles of ABA and GA during seed development 236
9.4.1.2 FUS3, a balancer of ABA and GA levels 236
9.4.1.3 AGL15, a transcriptional regulator of a GA deactivation gene 238
9.4.2 Regulation of ABA metabolism during seed imbibition in Arabidopsis 239
9.4.3 Regulation of GA metabolism during seed imbibition in Arabidopsis 240
9.4.3.1 Regulation of GA biosynthesis by light 240
9.4.3.2 Regulation of GA biosynthesis by cold temperature 241
9.5 Conclusions and perspectives 241
References 242
10 DE-repression of seed germination by GA signaling 248
CAMILLE M. STEBER
10.1 Introduction 248
10.2 Control of germination by GA signaling 248
10.3 The role of the ubiquitin鈥損roteasome pathway in GA signaling 252
10.4 Is RGL2 a 'master regulator' of seed germination? 255
10.5 Sleepyl is a positive regulator of seed germination in Arabidopsis 257
10.6 Do DELLA proteins have a conserved role in seed germination? 258
10.7 Future directions 260
References 260
11 Mechanisms and genes involved in germination sensu stricto 264
HIROYUKI NONOGAKI, FENG CHEN AND KENT J. BRADFORD
11.1 Introduction 264
11.2 Imbibition and water relations of seed germination 264
11.3 Testa/endosperm restraint and embryo growth potential 272
11.3.1 Testa and pericarp 272
11.3.2 Endosperm 273
11.3.3 Cell wall proteins and hydrolases involved in weakening of covering tissues 276
11.3.3.1 Expansins 276
11.3.3.2 Xyloglucan endotransglycosylase/hydrolases 277
11.3.3.3 Endo-β-mannanase, α-galactosidase, and β-mannosidase 279
11.3.3.4 Cellulase, arabinosidase, xylosidase 282
11.3.3.5 Polygalacturonase and pectin methylesterase 283
11.3.3.6 β-1,3-Glucanase and chitinase 283
11.3.3.7 Concerted action of cell wall hydrolases and expansins 285
11.3.4 Embryo growth potential 286
11.3.4.1 Generation of embryo growth potential 286
11.3.4.2 Gene expression associated with embryo growth 288
11.4 Approaches to identify additional genes involved in germination 289
11.4.1 Transcriptome and proteome analyses 289
11.4.2 Activation tagging and enhancer trapping 292
11.4.3 Potential involvement of microRNAs in seed germination 294
References 295
12 Sugar and abscisic acid regulation of germination and transition to seedling growth 305
BAS J.W. DEKKERS AND SJEF C.M. SMEEKENS
12.1 Introduction 305
12.2 ABA signaling during germination and early seedling growth 305
12.2.1 ABA response mutants isolated in germination-based screens 305
12.2.2 ABA inhibition of seed germination is suppressed by sugars 306
12.2.3 ABA blocks the transition from embryonic to vegetative growth 307
12.3 Sugar signaling represses germination and the transition to vegetative growth 309
12.3.1 Plant sugar signaling and the identification of sugar-response mutants 309
12.3.2 The glucose-insensitive response pathway 311
12.3.3 Other factors affecting the glucose response during early seedling development 314
12.3.4 Sugar delays seed germination in Arabidopsis 315
12.3.5 Imbibed seeds rapidly lose sensitivity for the glucose-induced germination delay 319
12.4 Conclusions 321
References 322
Index 329
List of Contributors xiii
Preface xv
1 Genetic control of seed development and seed mass 1
MASA-AKI OHTO, SANDRA L. STONE AND JOHN J. HARADA
1.1 Introduction 1
1.2 Overview of seed development in angiosperms 1
1.3 Genetic control of embryo development 3
1.3.1 Central regulators of embryogenesis 3
1.3.2 Genes involved in the morphogenesis phase of embryo development 4
1.3.3 Regulators of the maturation phase of embryo development 5
1.4 Genetic control of endosperm development 6
1.4.1 Genes required for cereal endosperm development 7
1.4.2 Genes that repress autonomous endosperm development 7
1.5 Genetic aspects of testa development 8
1.5.1 Genetic regulation of flavonoid biosynthesis and accumulation 9
1.5.2 Regulators of mucilage biosynthesis and accumulation 9
1.6 Control of seed mass 10
1.6.1 Genetic factors affecting seed mass 10
1.6.2 Testa development and seed mass 11
1.6.3 Endosperm development and seed mass 11
1.6.4 Sugar transport and metabolism during seed development 13
1.6.5 Metabolic control of seed development and size 15
1.7 Perspective 17
References 17
2 Seed coat development and dormancy 25
ISABELLE DEBEAUJON, LOIC LEPINIEC, LUCILLE POURCEL AND JEAN-MARC ROUTABOUL
2.1 Introduction 25
2.2 Development and anatomy of the seed coat 25
2.2.1 The seed envelopes 25
2.2.2 The Arabidopsis testa 27
2.3 Role of the seed coat in seed dormancy and germination 29
2.3.1 Constraints imposed by the seed coat 29
2.3.2 Flavonoids in Arabidopsis seeds 31
2.3.2.1 Main flavonoid end-products present in seeds 31
2.3.2.2 Molecular genetics of flavonoid metabolism 32
2.3.2.3 Effects of flavonoids on seed dormancy and germination 35
2.3.3 Flavonoids in seed dormancy and germination of various species 38
2.3.3.1 Solanaceae 38
2.3.3.2 Water permeability of testae in Leguminosae and other species 39
2.3.3.3 Flavonoids and other phenolics as direct and indirect germination inhibitors 39
2.3.3.4 Pre-harvest sprouting (PHS) in cereals 40
2.3.3.5 Heteromorphism and physiological heterogeneity among seeds 40
2.3.3.6 Interactions with endosperm 41
2.4 Link between seed coat-imposed dormancy and longevity 41
2.5 Concluding remarks 42
References 43
3 Definitions and hypotheses of seed dormancy 50
HENK W.M. HILHORST
3.1 Introduction 50
3.2 Classifications of dormancy 50
3.2.1 Endogenous dormancy 52
3.2.2 Exogenous dormancy 53
3.3 Definitions of dormancy 54
3.4 Primary dormancy 57
3.4.1 Induction of primary dormancy 57
3.4.1.1 Role of ABA in dormancy induction 57
3.4.1.2 Developmental programs and dormancy induction 58
3.4.2 Release of primary dormancy 59
3.4.2.1 After-ripening 59
3.4.2.2 Regulation of dormancy in imbibed seeds 60
3.5 Secondary dormancy 63
3.6 Signaling in dormancy 64
3.6.1 Stress signaling 64
3.6.2 Signaling networks 65
3.6.3 Environmental signals 65
3.7 Challenges for the future 67
References 67
4 Modeling of seed dormancy 72
PHIL S. ALLEN, ROBERTO L. BENECH-ARNOLD, DIEGO BATLLA AND KENT J. BRADFORD
4.1 Introduction 72
4.2 Types and phenology of seed dormancy 73
4.3 Environmental control of dormancy 76
4.3.1 Factors affecting dormancy levels of seed populations 76
4.3.1.1 Temperature 76
4.3.1.2 After-ripening 78
4.3.1.3 Stratification 79
4.3.2 Factors that stimulate germination 80
4.3.2.1 Fluctuating temperature 80
4.3.2.2 Light 81
4.3.2.3 Nitrate 82
4.3.3 Conceptual scheme of dormancy and its relationship to modeling 82
4.4 Approaches to modeling seed dormancy 82
4.4.1 Temperature response models and thermal time 83
4.4.2 Water potential responses and hydrotime models 87
4.4.3 Interactions of temperature and water potential 89
4.4.4 Modeling responses to other factors affecting dormancy and germination 90
4.5 Examples of seed dormancy models 91
4.5.1 Solanum tuberosum 91
4.5.2 Bromus tectorum 92
4.5.3 Polygonum aviculare 96
4.5.3.1 Modeling seed germination responses to temperature 97
4.5.3.2 Modeling seed responses to germination-stimulating factors 100
4.6 Population-based threshold models of seed dormancy 100
4.7 Conclusions and future directions 105
References 106
5 Genetic aspects of seed dormancy 113
LE脫NIE BENTSINK, WIM SOPPE AND MAARTEN KOORNNEEF
5.1 Introduction 113
5.2 Mutant approaches in Arabidopsis 114
5.3 Mutant approaches in other species 118
5.4 Genetic analyses of natural variation 119
5.4.1 Genetic analysis of natural variation in Arabidopsis 120
5.4.2 Natural variation for dormancy in grasses 120
5.5 What do the genetics teach us about dormancy and germination? 125
References 127
6 Lipid metabolism in seed dormancy 133
STEVEN PENFIELD, HELEN PINFIELD-WELLS AND IAN A. GRAHAM
6.1 Introduction 133
6.2 Metabolic pathways for TAG breakdown and conversion to sucrose 135
6.2.1 TAG hydrolysis and activation 135
6.2.2 Import of fatty acids into the peroxisome 135
6.2.3 Activation of fatty acids to acyl-CoA thioesters for β-Oxidation 136
6.2.4 β-Oxidation 137
6.2.4.1 Acyl-CoA oxidases 137
6.2.4.2 Multifunctional protein 137
6.2.4.3 3-L-Ketoacyl-CoA thiolase 138
6.2.4.4 Peroxisomal citrate synthase 138
6.2.5 Glyoxylate cycle and gluconeogenesis 138
6.2.5.1 Isocitrate lyase 139
6.2.5.2 Malate synthase 139
6.2.5.3 Phosphoenolpyruvate carboxylase 139
6.3 Lipid metabolism and seed dormancy 140
6.3.1 Importance of the ABC transporter for the transition from dormancy to germination 140
6.3.2 Defects in β-oxidation enzymes, but not in LACS, affect seed dormancy 142
6.3.3 Storage lipid mobilization (glyoxylate cycle and gluconeogenesis) is not required for seed dormancy release 144
6.4 Mechanisms for the involvement of β-oxidation in dormancy release 145
6.4.1 β-Oxidation does not fuel seed germination 145
6.4.2 β-Oxidation and hormonal signaling 145
6.4.3 Possible biosynthetic roles for β-oxidation in regulating germination 147
6.4.4 β-Oxidation, reactive oxygen species, and redox control 148
6.5 Conclusions 149
References 149
7 Nitric oxide in seed dormancy and germination 153
PAUL C. BETHKE, IGOR G.L. LIBOUREL AND RUSSELL L. JONES
7.1 Nitric oxide in plant growth and development 153
7.2 Challenges in NO chemistry and biology 153
7.3 Tools used in NO research 154
7.4 Roles of NO and other N-containing compounds in seed dormancy and germination 157
7.4.1 Nitrate, nitrite, and ammonium 158
7.4.2 Cyanide and azide 159
7.4.3 NO donors and germination 160
7.4.4 NO scavengers and germination 162
7.5 Biochemical and molecular basis of NO action in seeds 162
7.5.1 Synthesis of NO by plants 162
7.5.2 NO binding to metal-containing proteins 164
7.5.3 NO as an antioxidant 166
7.6 Interactions between NO and phytochrome or ABA 167
7.7 Ecological significance of NO 168
7.7.1 Nitrogen and vegetation gap sensing 168
7.7.2 Smoke and NO 169
7.8 Unresolved questions and concluding remarks 169
References 171
8 A merging of paths: abscisic acid and hormonal cross-talk in the control of seed dormancy maintenance and alleviation 176
J. ALLAN FEURTADO AND ALLISON R. KERMODE
8.1 Introduction 176
8.2 Abscisic acid 177
8.2.1 ABA in seed maturation and the induction of primary dormancy 177
8.2.2 Transcription factors and combinatorial control of seed development and maturation 180
8.2.3 ABA in dormancy maintenance and termination 182
8.2.3.1 ABA synthesis and homeostasis during dormancy maintenance and termination 182
8.2.3.2 ABA signaling factors and the control of dormancy maintenance and termination 186
8.3 Gibberellin 190
8.3.1 GA is antagonistic to ABA during seed development 190
8.3.2 GA promotes the transition to germination 191
8.4 Light interactions 195
8.4.1 GA synthesis and signaling are promoted by light through the action of phytochrome 195
8.4.2 ABA-associated signaling processes are opposed by light signaling 196
8.5 Ethylene 197
8.5.1 Ethylene counteracts ABA during seed development 197
8.5.2 Ethylene promotes the transition from dormancy to germination 198
8.6 Auxin and cytokinin 201
8.6.1 Auxin and cytokinin establish the embryo body plan during seed development 201
8.6.2 Auxin and cytokinins have not been intimately linked to dormancy maintenance or termination 202
8.7 Brassinosteroids 203
8.8 G-protein signaling reveals integration of GA, BR, ABA, and sugar responses 205
8.9 Profiling of hormone metabolic pathways in Arabidopsis mutants reveals cross-talk 206
8.10 Summary and future directions 208
References 211
9 Regulation of ABA and GA levels during seed development and germination in Arabidopsis 224
SHINJIRO YAMAGUCHI, YUJI KAMIYA AND EIJI NAMBARA
9.1 Introduction 224
9.2 Biosynthetic and deactivation pathways of ABA and GA 225
9.2.1 ABA biosynthesis 225
9.2.2 ABA deactivation 229
9.2.3 ABA-deficient mutants and seed germination 230
9.2.4 GA biosynthesis 230
9.2.5 GA deactivation 231
9.2.6 GA-deficient mutants and seed germination 231
9.3 Inhibitors of ABA and GA metabolism: efficacy and side effects of drugs 232
9.3.1 Drugs to reduce endogenous ABA levels 232
9.3.2 Drugs to increase endogenous ABA levels 234
9.3.3 Drugs to reduce GA levels 235
9.3.4 Side effects of drugs 235
9.4 Regulation of ABA and GA levels in Arabidopsis seeds 236
9.4.1 Regulation of ABA and GA levels during seed development 236
9.4.1.1 Roles of ABA and GA during seed development 236
9.4.1.2 FUS3, a balancer of ABA and GA levels 236
9.4.1.3 AGL15, a transcriptional regulator of a GA deactivation gene 238
9.4.2 Regulation of ABA metabolism during seed imbibition in Arabidopsis 239
9.4.3 Regulation of GA metabolism during seed imbibition in Arabidopsis 240
9.4.3.1 Regulation of GA biosynthesis by light 240
9.4.3.2 Regulation of GA biosynthesis by cold temperature 241
9.5 Conclusions and perspectives 241
References 242
10 DE-repression of seed germination by GA signaling 248
CAMILLE M. STEBER
10.1 Introduction 248
10.2 Control of germination by GA signaling 248
10.3 The role of the ubiquitin鈥損roteasome pathway in GA signaling 252
10.4 Is RGL2 a 'master regulator' of seed germination? 255
10.5 Sleepyl is a positive regulator of seed germination in Arabidopsis 257
10.6 Do DELLA proteins have a conserved role in seed germination? 258
10.7 Future directions 260
References 260
11 Mechanisms and genes involved in germination sensu stricto 264
HIROYUKI NONOGAKI, FENG CHEN AND KENT J. BRADFORD
11.1 Introduction 264
11.2 Imbibition and water relations of seed germination 264
11.3 Testa/endosperm restraint and embryo growth potential 272
11.3.1 Testa and pericarp 272
11.3.2 Endosperm 273
11.3.3 Cell wall proteins and hydrolases involved in weakening of covering tissues 276
11.3.3.1 Expansins 276
11.3.3.2 Xyloglucan endotransglycosylase/hydrolases 277
11.3.3.3 Endo-β-mannanase, α-galactosidase, and β-mannosidase 279
11.3.3.4 Cellulase, arabinosidase, xylosidase 282
11.3.3.5 Polygalacturonase and pectin methylesterase 283
11.3.3.6 β-1,3-Glucanase and chitinase 283
11.3.3.7 Concerted action of cell wall hydrolases and expansins 285
11.3.4 Embryo growth potential 286
11.3.4.1 Generation of embryo growth potential 286
11.3.4.2 Gene expression associated with embryo growth 288
11.4 Approaches to identify additional genes involved in germination 289
11.4.1 Transcriptome and proteome analyses 289
11.4.2 Activation tagging and enhancer trapping 292
11.4.3 Potential involvement of microRNAs in seed germination 294
References 295
12 Sugar and abscisic acid regulation of germination and transition to seedling growth 305
BAS J.W. DEKKERS AND SJEF C.M. SMEEKENS
12.1 Introduction 305
12.2 ABA signaling during germination and early seedling growth 305
12.2.1 ABA response mutants isolated in germination-based screens 305
12.2.2 ABA inhibition of seed germination is suppressed by sugars 306
12.2.3 ABA blocks the transition from embryonic to vegetative growth 307
12.3 Sugar signaling represses germination and the transition to vegetative growth 309
12.3.1 Plant sugar signaling and the identification of sugar-response mutants 309
12.3.2 The glucose-insensitive response pathway 311
12.3.3 Other factors affecting the glucose response during early seedling development 314
12.3.4 Sugar delays seed germination in Arabidopsis 315
12.3.5 Imbibed seeds rapidly lose sensitivity for the glucose-induced germination delay 319
12.4 Conclusions 321
References 322
Index 329
Seed development, dormancy and germination /
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