Annual Plant Reviews, The Plant Hormone Ethylene

Michael McManus is Professor at the Institute of Molecular Biosciences at Massey University, New Zealand. He is also an Editorial Board Member of Annual Plant Reviews.

• List of Contributors xv Preface xxiii1 100 Years of Ethylene – A Personal View 1Don Grierson1.1 Introduction 11.2 Ethylene biosynthesis 21.3 Ethylene perception and signalling 71.4 Differential responses to ethylene 91.5 Ethylene and development 101.6 Looking ahead 13Acknowledgements 14References 142 Early Events in the Ethylene Biosynthetic Pathway – Regulation of the Pools of Methionine and S-Adenosylmethionine 19Katharina B¨ urstenbinder and Margret Sauter2.1 Introduction 202.2 The metabolism of Met and SAM 222.3 Regulation of de novo Met synthesis 252.4 Regulation of the SAM pool 272.4.1 Regulation of SAMS genes by ethylene and of SAMS enzyme activity by protein-S-nitrosylation 292.5 The activated methyl cycle 302.6 The S-methylmethionine cycle 322.7 The methionine or Yang cycle 352.7.1 The Yang cycle in relation to polyamine and nicotianamine biosynthesis 392.7.2 Regulation of the Yang cycle in relation to ethylene synthesis 402.8 Conclusions 42Acknowledgement 43References 443 The Formation of ACC and Competition Between Polyamines and Ethylene for SAM 53Smadar Harpaz-Saad, Gyeong Mee Yoon, Autar K. Mattoo, and Joseph J. Kieber3.1 Introduction 533.2 Identification and characterization of ACC synthase activity in plants 543.2.1 Historical overview 543.2.2 Purification and properties of the ACC synthase protein 563.3 Analysis of ACC synthase at the transcriptional level 583.3.1 Molecular cloning of ACC synthase genes 583.3.2 Transcriptional regulation of the ACC synthase gene family 593.4 Post-transcriptional regulation of ACS 623.4.1 Identification and characterization of interactions with ETO1 623.4.2 Regulation of ACS degradation 643.5 Does ACC act as a signal? 653.6 Biosynthesis and physiology of polyamines 673.6.1 SAM is a substrate for polyamines 673.6.2 Physiology of polyamine effects in vitro and in vivo 673.6.3 Concurrent biosynthesis of ethylene and polyamines 703.6.4 Do plant cells invoke a homeostatic regulation of SAM levels? 72Acknowledgements 72References 724 The Fate of ACC in Higher Plants 83Sarah J. Dorling and Michael T. McManus4.1 Introduction 834.2 History of the discovery of ACC oxidase as the ethylene-forming enzyme 844.2.1 Early characterization of ACC oxidase 844.2.2 Cloning of the ethylene-forming enzyme as an indicator of enzyme activity 854.2.3 Initial biochemical demonstration of ethylene-forming enzyme activity in vitro 864.3 Mechanism of the ACC oxidase-catalyzed reaction 864.3.1 Investigation of the ACO reaction mechanism 874.3.2 Metabolism of HCN 894.3.3 Evidence of the conjugation of ACC 914.4 Transcriptional regulation of ACC oxidase 924.4.1 ACO multi-gene families 924.4.2 Differential expression of members of ACO multi-gene families in response to developmental and environmental stimuli 944.4.3 Transcriptional regulation of ACO gene expression 964.4.4 Crosstalk between ethylene signalling elements and ACO gene expression 974.5 Translational regulation of ACC oxidase 974.6 Evidence that ACC oxidase acts as a control point in ethylene biosynthesis 1004.6.1 Cell-specific expression of ACC oxidase 1024.6.2 Differential expression of ACS and ACO genes 1034.7 Evolutionary aspects of ACC oxidase 104Acknowledgements 105References 1055 Perception of Ethylene by Plants – Ethylene Receptors 117Brad M. Binder, Caren Chang and G. Eric Schaller5.1 Historical overview 1185.2 Subfamilies of ethylene receptors and their evolutionary history 1205.3 Ethylene binding 1235.3.1 Requirements for a metal cofactor 1235.3.2 Characterization of the ethylene-binding pocket and signal transduction 1245.4 Signal output from the receptors 1265.5 Overlapping and non-overlapping roles for the receptor isoforms in controlling various phenotypes 1285.6 Post-translational regulation of the receptors 1315.6.1 Clustering of receptors 1315.6.2 Ethylene-mediated degradation of receptors 1325.6.3 Regulatory role of REVERSION-TO-ETHYLENE SENSITIVITY1 (RTE1)/GREEN-RIPE (GR) 1335.6.4 Other proteins that interact with the ethylene receptors 1345.7 Conclusions and model 135Acknowledgements 137References 1386 Ethylene Signalling: the CTR1 Protein Kinase 147Silin Zhong and Caren Chang6.1 Introduction 1486.2 Discovery of CTR1, a negative regulator of ethylene signal transduction 1486.2.1 Isolation of the Arabidopsis CTR1 mutant 1486.2.2 CTR1 mutant phenotypes in Arabidopsis 1496.2.3 Placement of CTR1 in the ethylene-response pathway 1506.3 CTR1 Encodes a serine/threonine protein kinase 1516.3.1 Molecular cloning and sequence analysis of the Arabidopsis CTR1 gene 1516.3.2 CTR1 biochemical activity 1526.4 The CTR1 gene family 1536.4.1 The CTR multi-gene family in tomato 1536.4.2 Functional roles of tomato CTR genes 1536.4.3 Transcriptional regulation of CTR-like genes 1556.5 Regulation of CTR1 activity 1566.5.1 Physical association of CTR1 with ethylene receptors 1586.5.2 Membrane localization of CTR1 1596.5.3 An inhibitory role for the CTR1 N-terminus? 1596.5.4 Other factors that potentially interact with and regulate CTR1 activity 1606.6 Elusive targets of CTR1 signalling 1616.7 CTR1 crosstalk and interactions with other signals 1626.8 Conclusions 163Acknowledgements 164References 1647 EIN2 and EIN3 in Ethylene Signalling 169Young-Hee Cho, Sangho Lee and Sang-Dong Yoo7.1 Introduction 1697.2 Overview of ethylene signalling and EIN2 and EIN3 1727.3 Genetic identification and biochemical regulation of EIN2 1737.4 EIN3 regulation in ethylene signalling 1747.4.1 Genetic identification and biochemical regulation of EIN3 1747.4.2 Structural and functional analysis of ein3 function 1787.4.3 Function of EIN3 as transcription activator 1807.5 Functions of ERF1 and other ERFs in ethylene signalling 1817.6 Future directions 183Acknowledgements 184References 1848 Ethylene in Seed Development, Dormancy and Germination 189Renata Bogatek and Agnieszka Gniazdowska8.1 Introduction 1898.2 Ethylene in seed embryogenesis 1928.2.1 Ethylene biosynthesis during zygotic embryogenesis 1928.2.2 Ethylene involvement in the regulation of seed morphology 1948.3 Ethylene in seed dormancy and germination 1948.3.1 Ethylene biosynthesis during dormancy release and germination 1948.3.2 The role of ethylene in seed heterogeneity 1998.4 Ethylene interactions with other plant hormones in the regulation of seed dormancy and germination 1998.5 Ethylene interactions with ROS in the regulation of seed dormancy and germination 2028.6 Ethylene interactions with other small gaseous signalling molecules (NO, HCN) in the regulation of seed dormancy and germination 2048.7 Concluding remarks 207Acknowledgements 209References 2099 The Role of Ethylene in Plant Growth and Development 219Filip Vandenbussche and Dominique Van Der Straeten9.1 Introduction 2199.2 Design of root architecture 2209.3 Regulation of hypocotyl growth 2259.4 Shoot architecture and orientation: post-seedling growth 2299.4.1 Inhibition of growth by ethylene 2299.4.2 Stimulation of growth by ethylene 2299.4.3 Shoot gravitropism 2319.4.4 Control of stomatal density and aperture 2319.4.5 Activity of the shoot apical meristem 2319.5 Floral transition 2329.6 Determination of sexual forms of flowers 2329.7 Ethylene effects on growth controlling mechanisms 2339.8 Conclusions 234Acknowledgements 234References 23410 Ethylene and Cell Separation Processes 243Zinnia H. Gonzalez-Carranza and Jeremy A. Roberts10.1 Introduction 24310.2 Overview of the cell separation process 24410.2.1 Abscission 24510.2.2 Dehiscence 24910.2.3 Aerenchyma formation 25110.2.4 Stomata development and hydathode formation 25210.2.5 Root cap cell sloughing and lateral root emergence 25410.2.6 Xylem differentiation 25710.3 Transcription analyses during cell separation 25810.4 Relationship between ethylene and other hormones in the regulation of cell separation 25910.4.1 Ethyene and IAA 25910.4.2 Ethylene and jasmonic acid 26010.4.3 Ethylene and abscisic acid 26110.5 Ethylene and signalling systems during cell separation 26110.5.1 Role of IDA, IDA-like, HAESA and HAESA-like genes 26110.5.2 MAP kinases 26210.5.3 Nevershed 26210.6 Application of knowledge of abscission to crops of horticultural and agricultural importance 26210.7 Conclusions and future perspectives 263References 26511 Ethylene and Fruit Ripening 275Jean-Claude Pech, Eduardo Purgatto, Mondher Bouzayen and Alain Latch´e11.1 Introduction 27611.2 Regulation of ethylene production during ripening of climacteric fruit 27611.2.1 Regulation of ethylene biosynthesis genes during the System 1 to System 2 transition 27711.2.2 ACS gene alleles are major determinants of ethylene biosynthesis and shelf-life of climacteric fruit 28011.2.3 Genetic determinism of the climacteric character 28111.3 Transcriptional control of ethylene biosynthesis genes 28211.4 Role of ethylene in ripening of non-climacteric fruit 28311.5 Manipulation of ethylene biosynthesis and ripening 28411.6 Ethylene-dependent and -independent aspects of climacteric ripening 28611.7 Ethylene perception and transduction effects in fruit ripening 28811.7.1 Ethylene perception 28811.7.2 Chemical control of the post-harvest ethylene response in fruit ripening 28911.7.3 Ethylene signal transduction 29011.7.4 The transcriptional cascade leading to the regulation of ethylene-responsive and ripening-related genes 29111.8 Hormonal crosstalk in fruit ripening 29211.8.1 Ethylene and abscisic acid 29311.8.2 Ethylene and jasmonate 29311.8.3 Ethylene and auxin 29411.8.4 Ethylene and the gibberellins 29511.9 Conclusions and future directions 295Acknowledgements 296References 29612 Ethylene and Senescence Processes 305Laura E. Graham, Jos H.M. Schippers, Paul P. Dijkwel and Carol Wagstaff12.1 Introduction 30612.2 Overview of ethylene-mediated senescence in different plant organs 30612.2.1 Leaf senescence 30612.2.2 Pod senescence 31012.2.3 Petal senescence 31212.3 Transcriptional regulation of ethylene-mediated senescence processes 31412.3.1 Global regulation 31412.3.2 Transcription factors and signalling pathways 31512.4 Interaction of ethylene with other hormones in relation to senescence 32312.5 The importance of ethylene-mediated senescence in post-harvest biology 32512.5.1 Post-harvest factors affected by ethylene 32512.5.2 Ways of controlling ethylene-related post-harvest losses 32712.5.2.1 Packaging 32712.5.2.2 1-Methylcyclopropene 32812.6 Conclusions and future perspectives 329References 32913 Ethylene: Multi-Tasker in Plant–Attacker Interactions 343Sjoerd Van der Ent and Corn´e M.J. Pieterse13.1 Introduction 34413.2 Hormones in plant defence signalling 34613.2.1 Hormones as defence regulators 34613.2.2 Salicylic acid 34713.2.3 Jasmonic acid 34713.2.4 Ethylene 34813.3 Implications of ethylene in basal defence and disease susceptibility 34813.3.1 Studies with Arabidopsis thaliana 34813.3.2 Studies with tobacco 35013.3.3 Studies with tomato 35113.3.4 Studies with soybean 35213.3.5 Other plant species 35213.4 Implications of ethylene in systemic immune responses 35313.4.1 Systemic induced immunity 35313.4.2 Rhizobacteria-mediated ISR 35413.4.3 Genetic dissection of the ISR pathway in Arabidopsis 35613.4.4 Priming for enhanced JA/ethylene-dependent defences 35813.4.5 Molecular mechanisms of priming for enhanced defence 36013.4.6 Costs and benefits of priming for enhanced defence 36213.5 Ethylene modulates crosstalk among defence-signalling pathways 36213.5.1 Crosstalk in defence signalling 36213.5.2 Interplay among SA, JA and ethylene signalling 36313.5.3 Ethylene: an important modulator of defence-signalling pathways 36513.6 Concluding remarks 365Acknowledgements 366References 367Index 379First 8-page color plate section (between pages 168 and 169)Second 8-page color plate section (between pages 360 and 361)