Ethylene Action in Plants

Ethylene Action in Plants

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The discovery of the plant hormone ethylene was stunning—ethylene is a simple gas! Our expanding knowledge of the multiplicity of ethylene’s roles in plant development, physiology, and metabolism makes the study of this plant hormone increasingly compelling. Elucidation of the genetic regulation of ethylene biosynthesis, characterization of ethylene receptors and analysis of the pathway of ethylene signal transduction, coupled with the identifi- tion of components in the cascade and target genes, have provided insight into how this simple molecule can drive such a diversity of divergent processes. These scientific advances will lead to new technologies that will further enable researchers to harness the powers of ethylene for the benefit of agriculture. In Ethylene Action in Plants, classic and emerging roles of ethylene in plant developmental processes are integrated through recent advances ch- acterizing ethylene receptors, promoters and antagonists, and biological and environmental factors that mediate ethylene responses. The book’s editor, Dr. Nafees Khan, Aligarh Muslim University, Aligarh, India, an expert on ethylene with an impressive number of publications on the interactions between ethylene, photosynthesis, and growth of Brassica spp, brought together a highly qualified group of international experts to provide state-- the-art information. To simply list the topics included does not do justice to the book’s contents, as the articles are not just a compilation of the literature relevant to the topic.

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Published 10 May 2007
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Interaction of Ethylene and Other Compounds with the Ethylene Receptor: Agonists and Antagonists EDWARDC. SISLER, VARVARAP. GRICHKO ANDMARGRETHESEREK. . . . . . . . . . . . 1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2 Ethylene and Agonists. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.1 Discovery of Ethylene Action and Some Important Lessons from the Past . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.2 Molecular Requirements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2.3 Ethylene Binding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3 Ethylene Antagonists. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3.1 Chemicals Adjuvants Counteracting Ethylene . . . . . . . . . . . . . . . 1.3.2 Ethylene Agonists That Require Continuous Exposure to Give a Response. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3.3 Naturally Occurring Ethylene Antagonists . . . . . . . . . . . . . . . . . . 1.3.4 Photoactivated Compounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4 Ethylene Agonists That Require a Single Exposure . . . . . . . . . . . . . . . . 1.4.1 Ring Strain and Cyclopropene . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.2 1-Methylcyclopropene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.3 3-Methylcyclopropene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.4 3,3-Dimethylcyclopropene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.5 Other 1-Alkyl Cyclopropenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.6 Other Cyclopropenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.7 Is the Binding Site Restricted? . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.4.8 Hydrophobic Interaction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.5 How Does an Agonist Start a Signal? . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.6 Selected Papers on Significance of Ethylene Antagonists . . . . . . . . . . . 1.6.1 Application in Molecular Biology. . . . . . . . . . . . . . . . . . . . . . . . . 1.6.2 Application for Control of Plant Growth and Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.6.3 Commercial Use. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.6.4 Limitations of Compounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.6.5 Future Needs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.7 Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1 1 1
1 2 4 5 5
6 9 9 12 12 13 13 15 15 15 16 17 18 19 19
22 24 26 28 28 29
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Ethylene and Plant Growth DANNYTHOLEN, HENDRIKPOORTER ANDLAURENTIUSA.C.J. VOESENEK. . . . . . . . 2.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 Plant Growth Rate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2.1 Plant Growth Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Ethylene and Leaf Expansion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.4 Ethylene and Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.5 Ethylene and Growth under Optimal Conditions. . . . . . . . . . . . . . . . . . 2.6 Ethylene and Growth under Limiting Environmental Conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.7 Ethylene: Effects on Plant Growth? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ethylene and Leaf Senescence ANTONIOFERRANTE ANDALESSANDRAFRANCINI. . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Ethylene during Leaf Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 Ethylene-Induced Leaf Senescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3.1 Leaf Yellowing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3.2 Leaf Abscission. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 Effect of Ethylene Inhibitors, Promoters, and Ethylene-Releasing Compounds on Leaf Senescence . . . . . . . . . . . . . . . 3.5 Leaf Sensitivity and Ethylene Production . . . . . . . . . . . . . . . . . . . . . . . . 3.6 Effect of Ethylene on the Antioxidant System during Leaf Senescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.7 Ethylene and Other Plant Hormones during Leaf Senescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.8 Ethylene and Gene Expression during Leaf Senescence. . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Effect of Ethylene on Adventitious Root Formation JINXIANGWANG ANDRUICHIPAN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2 Ethylene Stimulates Adventitious Root Formation . . . . . . . . . . . . . . . . 4.3 Ethylene Inhibits Adventitious Rooting . . . . . . . . . . . . . . . . . . . . . . . . . 4.4 Ethylene Has No Role in Adventitious Root Formation . . . . . . . . . . . . 4.5 Interactions between Ethylene and Other Hormones . . . . . . . . . . . . . . 4.6 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.7 Outlook. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ethylene and Plant Responses to Abiotic Stress UWEDRUEGE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2 Abiotic Stress and Ethylene Biosynthesis . . . . . . . . . . . . . . . . . . . . . . . . 5.2.1 Wounding and Mechanical Stress . . . . . . . . . . . . . . . . . . . . . . . . 5.2.1.1 Wound Ethylene in Fruits. . . . . . . . . . . . . . . . . . . . . . . .
35 35 36 37 38 41 42
43 45 46
51 51 52 53 53 54
55 58
59
61 61 64
69 69 70 72 74 74 75 77 77
81 81 81 82 84
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5.2.1.2 Wound Ethylene in Vegetative Tissues. . . . . . . . . . . . . 5.2.1.3 Mechanical NonInjury Stress. . . . . . . . . . . . . . . . . . . . . 5.2.2 Water Deficit Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.3 Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.4 Flooding/Hypoxia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.5 Chilling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.6 Ozone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stress-Mediated Ethylene Sensitivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ethylene Action, Adaptation and Stress Tolerance . . . . . . . . . . . . . . . 5.4.1 Wound Response and Thigmomorphogenesis . . . . . . . . . . . . . 5.4.2 Ethylene and Ozone-Induced Cell Death . . . . . . . . . . . . . . . . . . 5.4.3 Ethylene and Root Stress-Mediated Shoot Growth. . . . . . . . . . 5.4.4 Ethylene and Adaptation to Salt Stress . . . . . . . . . . . . . . . . . . . 5.4.5 Ethylene and Flooding Tolerance . . . . . . . . . . . . . . . . . . . . . . . . 5.4.6 Ethylene and Chilling Response . . . . . . . . . . . . . . . . . . . . . . . . . Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ethylene in theRhizobiumLegume Symbiosis JEROENDENHERDER, SOFIEGOORMACHTIG ANDMARCELLEHOLSTERS. . . . . . . . 6.1 An Introduction to Legume Nodulation . . . . . . . . . . . . . . . . . . . . . . . . 6.2 Ethylene-Sensitive RHC Nodulation . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.1 Pharmacological Evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.2 Mutant Analysis and Transgenic Approaches in Plants and Bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.3 Ethylene Interferes with NF Signaling Within the Root Hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3 Ethylene is Indispensable for LRB Nodulation . . . . . . . . . . . . . . . . . . . 6.3.1 Pharmacological Data Show That Ethylene is Needed for Crack-Entry Invasion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.2 Ethylene Mediates the Phenotypic Plasticity in Root Nodule Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.3 Ethylene Mediates the Switch from Intercellular to Intracellular Invasion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.4 Ethylene Determines Nodule Primordium Positioning . . . . . . . . . . . . 6.5 Long-Distance Regulation Does Not Involve Ethylene . . . . . . . . . . . . 6.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
The Role of Ethylene in the Regulation of Stem Gravitropic Curvature MARCIAA. HARRISON. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1 The Modulating Role of Ethylene on Stem Gravitropic Curvature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2 Overview of Curvature Kinetics for Light-Grown Compared to Dark-Grown Tissues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3 Ethylene Production Increases after Horizontal Reorientation and during Curvature . . . . . . . . . . . . . . . . . . . . . . . . . . .
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86 88 89 91 93 95 97 98 102 103 103 106 107 109 109 110 111
119 119 121 121
123
125 125
126
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127 128 129 129 130
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Differing Sensitivities to Exogenous Ethylene may alter the Gravitropic Response . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4.1 Evidence of a Stimulatory Role for Ethylene in Stem Gravitropism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4.2 Ethylene’s Role in Slowing Gravitropic Curvature . . . . . . . . . . Ethylene and Auxin Cross-Talk in the Regulation of Gravitropic Curvature. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Role of Ethylene in Fruit Ripening PRAVENDRANATH, PRABODHK. TRIVEDI, VIDHUA. SANE AND ANIRUDHAP. SANE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.2 What is Fruit Ripening? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3 Ethylene Biosynthesis in Fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.1 ACC Synthase in Fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.2 ACC Oxidase in Fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.4 Ethylene Perception and Signal Transduction in Fruits . . . . . . . . . . . 8.5 Gene Expression and Regulation during Ripening in Fruits . . . . . . . . 8.5.1 Developmental Regulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.5.2 Ethylene Regualtion of Transcription Factors. . . . . . . . . . . . . . 8.5.3 Gene Expression during Softening . . . . . . . . . . . . . . . . . . . . . . . 8.5.4 Genes Involved in Changes in Pigments and Volatiles. . . . . . . 8.5.5 Other Ripening-Related Genes Isolated by Differential Screening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6 Roles of Other Hormones and Metabolites during Ripening . . . . . . . 8.6.1 Auxin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.2 Jasmonate and Jasmonic Acid . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.3 Abscisic Acid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.4 Other Metabolites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.7 Biotechnological Usage of Ethylene Biology . . . . . . . . . . . . . . . . . . . . . 8.7.1 Transgenic Fruits with Altered Ethylene Production . . . . . . . . 8.7.2 Transgenic Fruits with Altered Ethylene Perception . . . . . . . . 8.7.3 Transgenic Fruits with Altered Fruit Softening. . . . . . . . . . . . . 8.7.4 Ripening-Related Promoters. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ethylene Involvement in Photosynthesis and Growth NAFEESA. KHAN. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.2 Plant Growth Regulators and Photosynthetic Responses . . . . . . . . . . 9.3 Ethylene in Photosynthesis and Growth of Mustard (Brassica juncea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.3.1 Ethylene in Mustard Cultivars Differing In Photosynthetic Capacity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
140
140 143
144 147 148
151 151 152 154 154 155 156 158 159 159 160 162
162 163 164 164 165 166 167 167 170 171 174 176 176
185 185 185
187
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9.4
9.3.1.1 Effects of Ethylene Modulators. . . . . . . . . . . . . . . . . . . 9.3.1.2 Effects of ACC Synthase Activity Modulators. . . . . . . Ethylene in Photosynthesis and Growth of Defoliated Plants . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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