List of Contributors xv
Preface xxiii
1 100 Years of Ethylene ? A Personal View 1
Don Grierson
1.1 Introduction 1
1.2 Ethylene biosynthesis 2
1.3 Ethylene perception and signalling 7
1.4 Differential responses to ethylene 9
1.5 Ethylene and development 10
1.6 Looking ahead 13
Acknowledgements 14
References 14
2 Early Events in the Ethylene Biosynthetic Pathway ?
Regulation of the Pools of Methionine and
S-Adenosylmethionine 19
Katharina B¨ urstenbinder and Margret Sauter
2.1 Introduction 20
2.2 The metabolism of Met and SAM 22
2.3 Regulation of de novo Met synthesis 25
2.4 Regulation of the SAM pool 27
2.4.1 Regulation of SAMS genes by ethylene and of SAMS
enzyme activity by protein-S-nitrosylation 29
2.5 The activated methyl cycle 30
2.6 The S-methylmethionine cycle 32
2.7 The methionine or Yang cycle 35
2.7.1 The Yang cycle in relation to polyamine and nicotianamine
biosynthesis 39
2.7.2 Regulation of the Yang cycle in relation to ethylene
synthesis 40
2.8 Conclusions 42
Acknowledgement 43
References 44
3 The Formation of ACC and Competition Between Polyamines and
Ethylene for SAM 53
Smadar Harpaz-Saad, Gyeong Mee Yoon, Autar K. Mattoo, and
Joseph J. Kieber
3.1 Introduction 53
3.2 Identification and characterization of ACC synthase activity
in plants 54
3.2.1 Historical overview 54
3.2.2 Purification and properties of the ACC synthase protein
56
3.3 Analysis of ACC synthase at the transcriptional level 58
3.3.1 Molecular cloning of ACC synthase genes 58
3.3.2 Transcriptional regulation of the ACC synthase gene family
59
3.4 Post-transcriptional regulation of ACS 62
3.4.1 Identification and characterization of interactions with
ETO1 62
3.4.2 Regulation of ACS degradation 64
3.5 Does ACC act as a signal? 65
3.6 Biosynthesis and physiology of polyamines 67
3.6.1 SAM is a substrate for polyamines 67
3.6.2 Physiology of polyamine effects in vitro and in
vivo 67
3.6.3 Concurrent biosynthesis of ethylene and polyamines 70
3.6.4 Do plant cells invoke a homeostatic regulation of SAM
levels? 72
Acknowledgements 72
References 72
4 The Fate of ACC in Higher Plants 83
Sarah J. Dorling and Michael T. McManus
4.1 Introduction 83
4.2 History of the discovery of ACC oxidase as the
ethylene-forming enzyme 84
4.2.1 Early characterization of ACC oxidase 84
4.2.2 Cloning of the ethylene-forming enzyme as an indicator of
enzyme activity 85
4.2.3 Initial biochemical demonstration of ethylene-forming
enzyme activity in vitro 86
4.3 Mechanism of the ACC oxidase-catalyzed reaction 86
4.3.1 Investigation of the ACO reaction mechanism 87
4.3.2 Metabolism of HCN 89
4.3.3 Evidence of the conjugation of ACC 91
4.4 Transcriptional regulation of ACC oxidase 92
4.4.1 ACO multi-gene families 92
4.4.2 Differential expression of members of ACO multi-gene
families in response to developmental and environmental stimuli
94
4.4.3 Transcriptional regulation of ACO gene expression 96
4.4.4 Crosstalk between ethylene signalling elements and ACO
gene expression 97
4.5 Translational regulation of ACC oxidase 97
4.6 Evidence that ACC oxidase acts as a control point in
ethylene biosynthesis 100
4.6.1 Cell-specific expression of ACC oxidase 102
4.6.2 Differential expression of ACS and ACO genes 103
4.7 Evolutionary aspects of ACC oxidase 104
Acknowledgements 105
References 105
5 Perception of Ethylene by Plants ? Ethylene Receptors
117
Brad M. Binder, Caren Chang and G. Eric Schaller
5.1 Historical overview 118
5.2 Subfamilies of ethylene receptors and their evolutionary
history 120
5.3 Ethylene binding 123
5.3.1 Requirements for a metal cofactor 123
5.3.2 Characterization of the ethylene-binding pocket and signal
transduction 124
5.4 Signal output from the receptors 126
5.5 Overlapping and non-overlapping roles for the receptor
isoforms in controlling various phenotypes 128
5.6 Post-translational regulation of the receptors 131
5.6.1 Clustering of receptors 131
5.6.2 Ethylene-mediated degradation of receptors 132
5.6.3 Regulatory role of REVERSION-TO-ETHYLENE SENSITIVITY1
(RTE1)/GREEN-RIPE (GR) 133
5.6.4 Other proteins that interact with the ethylene receptors
134
5.7 Conclusions and model 135
Acknowledgements 137
References 138
6 Ethylene Signalling: the CTR1 Protein Kinase 147
Silin Zhong and Caren Chang
6.1 Introduction 148
6.2 Discovery of CTR1, a negative regulator of ethylene
signal transduction 148
6.2.1 Isolation of the Arabidopsis CTR1 mutant 148
6.2.2 CTR1 mutant phenotypes in Arabidopsis
149
6.2.3 Placement of CTR1 in the ethylene-response pathway
150
6.3 CTR1 Encodes a serine/threonine protein kinase
151
6.3.1 Molecular cloning and sequence analysis of the
Arabidopsis CTR1 gene 151
6.3.2 CTR1 biochemical activity 152
6.4 The CTR1 gene family 153
6.4.1 The CTR multi-gene family in tomato 153
6.4.2 Functional roles of tomato CTR genes 153
6.4.3 Transcriptional regulation of CTR-like genes
155
6.5 Regulation of CTR1 activity 156
6.5.1 Physical association of CTR1 with ethylene receptors
158
6.5.2 Membrane localization of CTR1 159
6.5.3 An inhibitory role for the CTR1 N-terminus? 159
6.5.4 Other factors that potentially interact with and regulate
CTR1 activity 160
6.6 Elusive targets of CTR1 signalling 161
6.7 CTR1 crosstalk and interactions with other signals 162
6.8 Conclusions 163
Acknowledgements 164
References 164
7 EIN2 and EIN3 in Ethylene Signalling 169
Young-Hee Cho, Sangho Lee and Sang-Dong Yoo
7.1 Introduction 169
7.2 Overview of ethylene signalling and EIN2 and EIN3 172
7.3 Genetic identification and biochemical regulation of EIN2
173
7.4 EIN3 regulation in ethylene signalling 174
7.4.1 Genetic identification and biochemical regulation of EIN3
174
7.4.2 Structural and functional analysis of ein3 function
178
7.4.3 Function of EIN3 as transcription activator 180
7.5 Functions of ERF1 and other ERFs in ethylene signalling
181
7.6 Future directions 183
Acknowledgements 184
References 184
8 Ethylene in Seed Development, Dormancy and Germination
189
Renata Bogatek and Agnieszka Gniazdowska
8.1 Introduction 189
8.2 Ethylene in seed embryogenesis 192
8.2.1 Ethylene biosynthesis during zygotic embryogenesis 192
8.2.2 Ethylene involvement in the regulation of seed morphology
194
8.3 Ethylene in seed dormancy and germination 194
8.3.1 Ethylene biosynthesis during dormancy release and
germination 194
8.3.2 The role of ethylene in seed heterogeneity 199
8.4 Ethylene interactions with other plant hormones in the
regulation of seed dormancy and germination 199
8.5 Ethylene interactions with ROS in the regulation of seed
dormancy and germination 202
8.6 Ethylene interactions with other small gaseous signalling
molecules (NO, HCN) in the regulation of seed dormancy and
germination 204
8.7 Concluding remarks 207
Acknowledgements 209
References 209
9 The Role of Ethylene in Plant Growth and Development
219
Filip Vandenbussche and Dominique Van Der Straeten
9.1 Introduction 219
9.2 Design of root architecture 220
9.3 Regulation of hypocotyl growth 225
9.4 Shoot architecture and orientation: post-seedling growth
229
9.4.1 Inhibition of growth by ethylene 229
9.4.2 Stimulation of growth by ethylene 229
9.4.3 Shoot gravitropism 231
9.4.4 Control of stomatal density and aperture 231
9.4.5 Activity of the shoot apical meristem 231
9.5 Floral transition 232
9.6 Determination of sexual forms of flowers 232
9.7 Ethylene effects on growth controlling mechanisms 233
9.8 Conclusions 234
Acknowledgements 234
References 234
10 Ethylene and Cell Separation Processes 243
Zinnia H. Gonzalez-Carranza and Jeremy A. Roberts
10.1 Introduction 243
10.2 Overview of the cell separation process 244
10.2.1 Abscission 245
10.2.2 Dehiscence 249
10.2.3 Aerenchyma formation 251
10.2.4 Stomata development and hydathode formation 252
10.2.5 Root cap cell sloughing and lateral root emergence
254
10.2.6 Xylem differentiation 257
10.3 Transcription analyses during cell separation 258
10.4 Relationship between ethylene and other hormones in the
regulation of cell separation 259
10.4.1 Ethyene and IAA 259
10.4.2 Ethylene and jasmonic acid 260
10.4.3 Ethylene and abscisic acid 261
10.5 Ethylene and signalling systems during cell separation
261
10.5.1 Role of IDA, IDA-like, HAESA and HAESA-like genes 261
10.5.2 MAP kinases 262
10.5.3 Nevershed 262
10.6 Application of knowledge of abscission to crops of
horticultural and agricultural importance 262
10.7 Conclusions and future perspectives 263
References 265
11 Ethylene and Fruit Ripening 275
Jean-Claude Pech, Eduardo Purgatto, Mondher Bouzayen and
Alain Latch´e
11.1 Introduction 276
11.2 Regulation of ethylene production during ripening of
climacteric fruit 276
11.2.1 Regulation of ethylene biosynthesis genes during the
System 1 to System 2 transition 277
11.2.2 ACS gene alleles are major determinants of ethylene
biosynthesis and shelf-life of climacteric fruit 280
11.2.3 Genetic determinism of the climacteric character 281
11.3 Transcriptional control of ethylene biosynthesis genes
282
11.4 Role of ethylene in ripening of non-climacteric fruit
283
11.5 Manipulation of ethylene biosynthesis and ripening 284
11.6 Ethylene-dependent and -independent aspects of climacteric
ripening 286
11.7 Ethylene perception and transduction effects in fruit
ripening 288
11.7.1 Ethylene perception 288
11.7.2 Chemical control of the post-harvest ethylene response in
fruit ripening 289
11.7.3 Ethylene signal transduction 290
11.7.4 The transcriptional cascade leading to the regulation of
ethylene-responsive and ripening-related genes 291
11.8 Hormonal crosstalk in fruit ripening 292
11.8.1 Ethylene and abscisic acid 293
11.8.2 Ethylene and jasmonate 293
11.8.3 Ethylene and auxin 294
11.8.4 Ethylene and the gibberellins 295
11.9 Conclusions and future directions 295
Acknowledgements 296
References 296
12 Ethylene and Senescence Processes 305
Laura E. Graham, Jos H.M. Schippers, Paul P. Dijkwel and
Carol Wagstaff
12.1 Introduction 306
12.2 Overview of ethylene-mediated senescence in different plant
organs 306
12.2.1 Leaf senescence 306
12.2.2 Pod senescence 310
12.2.3 Petal senescence 312
12.3 Transcriptional regulation of ethylene-mediated senescence
processes 314
12.3.1 Global regulation 314
12.3.2 Transcription factors and signalling pathways 315
12.4 Interaction of ethylene with other hormones in relation to
senescence 323
12.5 The importance of ethylene-mediated senescence in
post-harvest biology 325
12.5.1 Post-harvest factors affected by ethylene 325
12.5.2 Ways of controlling ethylene-related post-harvest losses
327
12.5.2.1 Packaging 327
12.5.2.2 1-Methylcyclopropene 328
12.6 Conclusions and future perspectives 329
References 329
13 Ethylene: Multi-Tasker in Plant?Attacker
Interactions 343
Sjoerd Van der Ent and Corn´e M.J. Pieterse
13.1 Introduction 344
13.2 Hormones in plant defence signalling 346
13.2.1 Hormones as defence regulators 346
13.2.2 Salicylic acid 347
13.2.3 Jasmonic acid 347
13.2.4 Ethylene 348
13.3 Implications of ethylene in basal defence and disease
susceptibility 348
13.3.1 Studies with Arabidopsis thaliana 348
13.3.2 Studies with tobacco 350
13.3.3 Studies with tomato 351
13.3.4 Studies with soybean 352
13.3.5 Other plant species 352
13.4 Implications of ethylene in systemic immune responses
353
13.4.1 Systemic induced immunity 353
13.4.2 Rhizobacteria-mediated ISR 354
13.4.3 Genetic dissection of the ISR pathway in
Arabidopsis 356
13.4.4 Priming for enhanced JA/ethylene-dependent defences
358
13.4.5 Molecular mechanisms of priming for enhanced defence
360
13.4.6 Costs and benefits of priming for enhanced defence
362
13.5 Ethylene modulates crosstalk among defence-signalling
pathways 362
13.5.1 Crosstalk in defence signalling 362
13.5.2 Interplay among SA, JA and ethylene signalling 363
13.5.3 Ethylene: an important modulator of defence-signalling
pathways 365
13.6 Concluding remarks 365
Acknowledgements 366
References 367
Index 379
First 8-page color plate section (between pages 168 and 169)
Second 8-page color plate section (between pages 360 and
361)
