Ethylene: Agricultural Sources and Applications

349 Pages·2002·12.92 MB·English

by  Muhammad Arshad

ETHYLENE
Agricultural Sources
and Applications
ETHYLENE
Agricultural Sources
and Applications
Muhammad Arshad
University of Agriculture
Faisalabad, Pakistan
and
William T. Frankenberger, Jr .
University of California
Riverside, California
Springer Science+Business Media, LLC
Library of Congress Cataloging-in-Publication Data
~had,~uhanunad, 1956-
Ethylene: agricultura! resources and applicationslby ~uharnmad Arshad and William
T. Frakenberger, Ir.
p. cm.
Inc1udes bibliographica1 references (p. ).
ISBN 978-1-4613-5189-4 ISBN 978-1-4615-0675-1 (eBook)
DOI 10.1007/978-1-4615-0675-1
1. Ethylene. 1. Frakenberger, W. T. (William T.), 1952- II. Title.
QK898.E8 A 78 2002
631.8'9-dc21
2001041356
ISBN 978-1-4613-5189-4
©2oo2 Springer Science+Business Media New York
Originally published by Kluwer Academic/Plenum Publishers, New York in 2002
Softcover reprint of the hardcover 1 st edition 2002
http://www.wkap.nl/
10987654321
A C.1.P. record for this book is available from the Library of Congress
AII rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any
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permission from the Publisher
PREFACE
With an ever-increasing demand for more food supply, agricultural scientists will
have to search for new ways and technologies to promote food production. In recent
decades, plant growth regulators (PGRs) have made great strides in promoting plant
growth and development. PGRs are organic compounds which have the ability to
dramatically affect physiological plant processes when present in extremely low
concentrations (in the range of micro- to picograms). Although all higher plants have the
ability to synthesize PGRs endogenously, they do respond to the exogenous sources most
likely due to not having the capacity to synthesize sufficient endogenous phytohormones
for optimal growth and development under given climatic and environmental conditions.
In recent years, PGRs have established their position as a new generation of
agrochemicals after pesticides, insecticides and herbicides. Interest in the commercial
use of PGRs for improving plant growth and crop yields is also increasing because of
their non-polluting nature. The use of PGRs in the post-harvest technology is well
established and many new breakthroughs have recently been revealed.
Ethylene (C2~) is one of the five major classes of PGRs. Although C2~ is a
simple gas, it has a very strong multifaceted role in plant physiology. Initially, known as
the "ripening hormone", C2~ is now considered to be involved in numerous processes
from seed ontogeny to senescence and even in post-harvest technology. Results of
several studies warrant that C2~ of exogenous sources could be used for the betterment
of agricultural production. The exogenous sources include natural production by soil
microbiota and synthetic C2~ releasing compounds. A commercial product "ethephon"
has already provided a breakthrough discovery against lodging in cereals. The
importance of C2~ as a potent plant growth regulator has led us to compile the most
recent scientific literature into this book.
The biochemistry of endogenous C2~ biosynthesis and its role in physiology of
higher plants have been compiled in the form of several books and reviews. However,
very little attention has been given to the soil microbiota and their potential exogenous
source of C2~' The ecological importance of microbially produced C2~ was evident
after the dramatic effects on plant growth were published in the 1990s. We have pooled
this information into a comprehensive book. In this book, we provide eight chapters
including an introduction of C2~ as a hormone; its role in plant physiology; sources and
factors affecting microbial production of C2H4 ; biochemistry of C2~ production by
microorganisms; production in soil; the role of C2~ in symbiosis and pathogenesis; and
its commercial application in agriculture. Overall, this book is extremely valuable in that
it covers all the aspects of C2H4 sources, in-depth biochemistry and applications.
Our comprehensive effort in providing this resource will benefit agricultural
scientists including physiologists, microbiologists, soil scientists, botanists, ecologists,
and horticulturalists to develop new technologies based on the release of C2~ for the
betterment of agricultural production.
Special acknowledgement is given to Louise DeHayes for typing this entire book and
to Linda Bobbitt for her assistance in the illustrations of the figures. We also wish to
thank Ben Johanson and Tariq Siddique for their assistance in proofreading the text.
v
CONTENTS
1. THE PLANT HORMONE, ETHYLENE ............................ 1
1.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2. Ethylene as a Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.3. Biological Activity of Ethylene ............ . . . . . . . . . . . . . . . . . . . . . . 2
1.4. Historical Perspective. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.5. Sources of Ethylene and Applications. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.6. Future Prospects .............................................. 6
1. 7. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2. ETHYLENE IN PLANT PHYSIOLOGY ..... . . . . . . . . . . . . . . . . . . . . . . . 11
2.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.2. Endogenous Production of Ethylene .............................. 11
2.3. Biosynthetic Pathway. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
2.3.1. Establishment of Methionine as a Physiological Precursor. . . . . . . 13
2.3.2. Conversion of Methionine into S-Adenosymethionine (Step I) ... 13
2.3.2.1. Biochemical and Molecular Properties of Methionine
Adenotransferase ................................. 14
2.3.3. Conversion of S-adenosymethionine Conversion into
1-Aminocyclopropane-l-carboxylic Acid (Step II) ............. 14
2.3.3.1. Biochemical and Molecular Properties of ACC Synthase 16
2.3.4. Conversion of l-Aminocyclopropane-l-carboxylic Acid into
Ethylene (Step III) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.3.4.1. Biochemical and Molecular Properties of ACC Oxidase 20
2.3.5. Conjugation of 1-Aminocyclopropane-l-carboxylic Acid. . . . . . . . 23
2.3.5.1. ACC N-Malonyltransferase . . . . . . . . . . . . . . . . . . . . . . . . . 24
2.3.6. Methionine Cycle ....................................... 25
2.4. Regulation of Ethylene Production ............................... 25
2.5. Metabolism. . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . .. . . . .. . . .. . . . . . . . . 27
2.6. Mechanism of Action and Ethylene Receptors . . . . . . . . . . . . . . . . . . . . . . 28
2.7. Ethylene Production by Lower Plants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
vii
viii Contents
2.8. Concluding Remarks .......................................... 38
2.9. References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
3. BIOCHEMISTRY OF MICROBIAL PRODUCTION OF ETHYLENE .. 51
3.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
3.2. SubstrateslPrecursors/Stimulators of Ethylene ...................... 51
3.3. Biosynthetic Pathways ......................................... 56
3.3.1. ACC Deaminase ........................................ 60
3.4. Methionine-Dependent and -Independent Pathways. . . . . . . . . . . . . . . . . . 64
3.4.1. Methionine-Dependent Pathway ................ . . . . . . . . . . . . 64
3.4.1.1. Biosynthesis of C H by Escherichia coli ... ,......... 64
2 4
3.4.1.2. Biosynthesis of C H by Cryptococcus albidus . . . . . . . . . 66
2 4
3.4.1.3. Biosynthesis of C H by Septoria musiva ............. 69
2 4
3.4.1.4. Summary ....................................... 70
3.4.2. Methionine-Independent Pathway . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
3.4.2.1. Biosynthesis of C H by Penicillium digitatum . . . . . . . . . 70
2 4
3.4.2.2. Biosynthesis of C H by Pseudomonas syringae pv.
2 4
phaseolicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
3.4.2.3. Biosynthesis of C H by Other Fungi ................ 80
2 4
3.4.2.4. Summary ....................................... 81
3.5. Molecular Aspects of the Ethylene-forming Enzyme of Pseudomonas
syringae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
3.6. Structure of Ethylene-forming Enzymes ........................... 83
3.7. Microbial Metabolism of Ethylene ............................... 85
3.8. Concluding Remarks .......................................... 90
3.9. References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
4. FACTORS AFFECTING MICROBIAL PRODUCTION OF ETHYLENE
97
4.1. Microbial Production of Ethylene ................................ 97
4.2. Factors Affecting Microbial Production of Ethylene in Vitro .......... 98
4.2.1. Carbon Sources ......................................... 104
4.2.2. Nitrogen and Phosphorus ................................. 105
4.2.3. pH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
4.2.4. Aeration ............................................... 118
4.2.5. Temperature... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 118
4.2.6. Trace Elements ......................................... 124
4.2.7. Growth Phase. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
4.3. Effect of Ethylene on Microbial Growth .......................... 128
4.3.1. Bacteria ............................................... 129
4.3.2. Fungi ................................................. 129
4.3.3. Soil Fungistasis ......................................... 132
4.4. Concluding Remarks .......................................... 132
4.5. References.................. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
CONTENTS ix
5. ETHYLENE IN SOIL 139
5.1. Ethylene Production in Soil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
5.1.1. Abiotic Production. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
5.1.2. Biotic Production. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
5.1.2.1. Types of Soil Microorganisms Involved ............. 143
5.2. Physicochemical Properties of Soil and Ethylene Production ......... 145
5.2.1. Soil Organic Matter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
5.2.2. Soil Aeration .......................................... 147
5.2.3. Soil Reaction .......................................... 154
5.2.4. Soil Temperature ....................................... 155
5.2.5. Soil Texture ........................................... 157
5.2.6. Soil Depth ............................................ 158
5.3. Organic Amendments and Ethylene Accumulation ................. 158
5.3.1. Organic Amendments with Defined Chemical Composition. . ... 158
5.3.2. Organic Amendments without Defined Chemical Composition .. 160
5.4. Inorganic Amendments and Ethylene Accumulation in Soil .......... 164
5.5. Nitrate as a Suppressor of Ethylene Accumulation in Soil ........... 169
5.6. Ferrous Iron as a Stimulator of C H Production in Soil. . . . . . . . . . . . . 172
2 4
5.7. Persistence of Ethylene in Soil ................................. 178
5.8. Plant Responses Evoked by Ethylene and Waterlogging ............. 183
5.9. Concluding Remarks ......................................... 189
5.10. References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
6. ETHYLENE IN SYMBIOSIS .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
6.1. Introduction................................................. 195
6.2. Symbiotic Associations ....................................... 195
6.3. Nodulation ................................................. 195
6.3.1. Effects of Exogenous Ethylene on Nodulation ............... 196
6.3.2. Effects of Exogenous Ethylene on Nod Factor(s) ............. 199
6.3.3. Accelerated Ethylene Evolution from Rhizobium-Infected Roots 202
6.3.4. Nitrate and Light-Induced Ethylene Production and Nodulation 206
6.3.5. Ethylene Sensitive and Insensitive Legume Mutants and
Nodulation ............................................ 207
6.3.6. Effects of Inhibitors of Ethylene Action or Biosynthesis on
Nodulation ............................................ 210
6.3.7. Summary ............................................. 214
6.4. Mycorrhizal Associations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 216
6.4.1. Ethylene Production by Mycosymbiont ..................... 217
6.4.2. Ethylene Production by Mycorrhizal Roots .................. 217
6.4.3. Exogenous Ethylene and Mycorrhizae ...................... 220
6.4.4. Relationship between in Vitro Ethylene Production and Growth
Response of Mycorrhizal Plants ........................... 222
6.4.5. Summary ............................................. 227
6.5. Lichen (An Algal-Fungal Symbiosis) ............................ 227
x CONTENTS
6.5.1. Ethylene Production by Lichens .. . . . . . . . . . . . . . . . . . . . . . . . . . . 227
6.5.2. Sources of Ethylene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
6.5.3. Biochemistry of Ethylene Production by Lichens .............. 231
6.5.4. Summary .............................................. 234
6.6. Concluding Remarks .......................................... 234
6.7. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235
7. ETHYLENE IN PATHOGENESIS ................................. 241
7.1. Infection: A Biotic Stress. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
7.2. Enhanced Ethylene Production during Pathogenesis ................. 241
7.3. Source of Accelerated Ethylene Evolution during Pathogenesis ........ 249
7.3.1. Contribution by the Pathogen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
7.3.2. Contribution by the Host. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 251
7.4. Role of Ethylene in Disease Development ......................... 254
7.4.1. Effect of Exogenous Ethylene ............................. 255
7.4.2. Ethylene Production and Virulence ......................... 256
7.4.3. Ethylene Production and Development of Disease Symptoms .... 258
7.4.4. Ethylene and Toxins Produced by Pathogens ................. 260
7.4.5. Ethylene Mutants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 261
7.5. Role of Ethylene in Disease Resistance ........................... 262
7.5.1. Induction of Pathogenesis Related (PR) Proteins and Enzymes ... 264
7.5.1.1. Chitinases and ~-1,3-Glucanases .................... 265
7.5.1.2. Phenylalanine Ammonia-Lyase. .. ... . .. . .. . . . . . .. ... 267
7.5.1.3. Ethylene Biosynthesis-Inducing Xylanase ............. 268
7.5.1.4. Others.......................................... 268
7.5.2. Induction of Phytoalexins and Antibiotics. . . . . . . . . . . . . . . . . . .. 269
7.5.3. Induction of Structural and Development Changes. . . . . . . . . . . . . 270
7.5.4. Summary .............................................. 271
7.6. Soil Ethylene as a Weedicide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 272
7.7. Concluding Remarks .......................................... 273
7.8. References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 275
8. ETHYLENE IN AGRICULTURE: SYNTHETIC AND NATURAL
SOURCES AND APPLICATIONS ................................. 289
8.1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
8.2. Ethylene Sources and Their Applications in Agriculture .. . . . . . . . . . . .. 290
8.2.1. Ethylene Gas ........................................... 290
8.2.2. Ethylene-Releasing Compounds.. . . . . . . . . .. . .. . .. . . . . . .. . .. 291
8.2.3. Ethephon and Its Applications ............................. 292
8.2.4. Retprol and Its Applications. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
8.2.5. Soil Ethylene and Its Ecological Significance ................. 305
8.3. Alterations in Endogenous Ethylene Biosynthesis and Plant Responses.. 315
8.3.1. Transgenic Plants with ACC Oxidase ACC Synthease Antisense 315
8.3.2. Transgenic Plants with Bacterial Ethylene-Forming Enzymes .... 317
CONTENTS xi
8.3.3. Transgenic Plants with Bacterial ACC-Deaminase ............. 318
8.3.4. Inoculation with ACC-Deaminase Containing Plant Growth-
Promoting Rhizobacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322
8.4. Concluding Remarks .......................................... 324
8.5. References..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327
INDEX ........................................................... 337
To my wife Talat, son Haaris
and daughters Habiba and Fatima
Muhammad Arshad
To my wife, Margaret E. Beebe-Frankenberger
William T. Frankenberger