Natural Growth Inhibitors and Phytohormones in Plants and Environment

Transcription

1 Natural Growth Inhibitors and Phytohormones in Plants and Environment

2 Natural Growth Inhibitors and Phytohormones in Plants and Environment by Valentine I. Kefeli Biomost, Inc., Pennsylvania, U.S.A. and Maria V. Kalevitch Robert Morris University, Pennsylvania, U.S.A. Editor Bruno Bors'ari Slippery Rock University, Pennsylvania, U.S.A. Springer-Science+Business Media, B.Y.

3 A C.LP. Catalogue record for this book is available from the Library of Congress. Printed on acid-free paper ISBN ISBN (ebook) DOI / All Rights Reserved 2003 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in Softcover reprint of the hardcover I st edition 2003 No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work.

4 CONTENTS PREFACE... V ACKNOWLEDGMENT... VII INTRODUCTION... IX CHAPTER SYSTEM OF GROWTH AND DEVELOPMENT REGULATION IN THE PLANT PROPERTIES OF NATURAL HORMONES Auxins... I Gibberellins Cytokinins PROPERTIES OF NATURAL INHIBITORS Ethylene Abscisic acid and phenolic inhibitors Biosynthesis of Phytohormones and Natural Growth Inhibitors CHAPTER NATURAL GROWTH INHIBITORS AND PHYTOHORMONES UNDER THE CONTROL OF THE PLANT'S GENOME GENOME - GROWTH REGULATORS PHOTOSYNTHETIC CHANGES DURING GENOME MUTATION PHOTOMORPHOGENESIS AND LIGHT EFFECT ON PHYTOHORMONES LiGHT AND FUNCTIONS OF PHENOLIC COMPOUNDS IN PHOTOMORPHOGENESIS LIGHT AND HORMONES NATURAL GROWTH REGULATORS AND FUNCTIONS OF CHLOROPLASTS CHAPTER NATURAL INHIBITORS AND PHYTOHORMONES DURING LEAVES GROWTH AND DEVELOPMENT CHLOROPHYLL BIOSYNTHESIS IN THE GREEN CHLOROPLASTS PHYTOHORMONES AND INHIBITORS IN MUTANTS PHENOLICS IN CHLOROPLASTS CHAPTER NATURAL GROWTH INHIBITORS AND PHYTOHORMONES IN THE PROCESS OF CELLS AND ORGANS ELONGATION I

5 II 4.1 THE ROLE OF PHYTOHORMONES AND NATURAL INHIBITORS IN GROWTH OF ABOVEGROUND ORGANS OF PLANTS CHAPTER PHENOLIC INHIBITORS AND ABSCISIC ACID DURING DORMANCY NATURAL INHIBITORS, AUXINS AND PLANT'S DORMANCy ROLE OF NATURAL GROWTH INHIBITORS DURING PLANT'S DORMANCY FUNCTIONS OF PHYTOHORMONES AND GROWTH INHIBITORS DURING DORMANCY TERMINATION NATURAL GROWTH INHIBITORS AND SEEDS GERMINATION CHAPTER CATABOLISM OF NATURAL GROWTH INHIBITORS IN THE PLANT AND IN THE ABSCISED LEAVES TRANSFORMATIONS OF NATURAL GROWTH INHIBITORS DYNAMICS OF DECOMPOSITION OF ISOSALIPURPOSIDE ARID PHLORIDZIN BY PLANT TISSUES DECOMPOSITION PRODUCTS OF ISOSALIPURPOSIDE AND PHLORIDZIN BIOLOGICAL ACTIVITY OF PHLORIDZIN AND ITS AGLYCONE PHLORETIN ETHYLENE EVOLUTION AND ABSCISIC ACID CONTENT IN CITRUS PLANTS INTACT AND ABSCISED LEAVES AND THEIR PROPERTIES CHAPTER NATURAL GROWTH INHIBITORS AS ALLELOPATHOGENS AND BOTANICAL HERBICIDES CHAPTER NATURAL GROWTH INHIBITORS AND STRESS CONDITIONS GENERAL ASPECTS OF STRESS EFFECTS Low TEMPERATURE EFFECTS LIGHT AS A FACTOR OF ONTOGENY AND STRESS Exogenous and endogenous systems of photo morphogenesis and phototropism in plants Photoreceptors Light and phototropism nhibiting effects of light intensity Plant Height Surface Area of Leaflets and Stipules Specific Leaf Weight (SLW) Daily Dry Weight Gain Growth regulators and light deficiency UV-light as stress factor

6 Light and ethylene production UV-B light and ABA-ethylene ratio CHAPTER NATURAL GROWTH INHIBITORS AND PHYTOHORMONES IN THE INTACT PLANTS AND ISOLATED CELLS, ORGANS AND TISSUES CALLUS PRODUCTION BY WILD-TYPE AND TRANSGENIC REGENERANTS PROLIFERATION OF SUBCULTURED CALLI CHAPTER NATURAL GROWTH INHIBITORS AND BIOTESTS ACTION OF METABOLIC INHIBITORS AND ANTIBIOTICS ON GROWTH IN BIO- ASSAYS COMPARISON OF THE EFFECTS OF NATURAL GROWTH INHIBITORS, ANTIBIOTICS AND SOME SOLVENTS ON GROWTH OF COLEOPTILE SEGMENTS AND ROOT FORMATION IN PHASEOLUS CUTTINGS COMPARISON OF THE EFFECTS OF NATURAL AND SYNTHETIC GROWTH INHIBITORS ON VARIOUS GROWTH FORMS EFFECT OF NATURAL INHIBITORS ON PHYTOHORMONE ACTIVITY AND ON SOME METABOLIC PROCESSES IN PLANTS BIO-ASSAYMETHODS Cell expansion tests Cell division tests Complex tests ANALYSIS OF INTERACTION OF GROWTH REGULATORS CONCLUSION REFERENCES GLOSSARY INDEX III

7 PREFACE This book represents the authors' lifetime dedication to the study of inhibitors and phytohormones as well as its practical applications for achieving a more sustainable agriculture. Their work focuses on the functions of various groups of active molecules, their direct effect upon plant growth, but also implications for their impact upon the surrounding environment are explored. The main idea of the book evolved from the need to determine a balance among natural growth inhibitors and phytohormones. This approach was pursued through a better understanding of their biochemical pathways, their effects on plants physiological functions, and their influence upon stress factors on plant ontogenesis. Therefore, this effort proposes a more holistic approach to the study of plant physiology, in which the plant-soil interactions are discussed, with a profound description of different allelochemicals and their effects on plants growth. A rigorous attention is also paid to discuss the role of microorganisms in ecosystems and their capability to synthesize physiologically active substances, which trigger also unique plant-microbial interactions. These synergies are leading scientists to the discovery of major breakthroughs in agriculture and pharmacology that are revolutionizing old epistemologies and thus, contributing to the emergence of a philosophy of interconnectedness for the whole biosphere. The initial conceptions about plant hormone-inhibitor interactions were based upon the research accomplished by the authors at the Russian Academy of Sciences during the last three decades. The outcome of these efforts have been published in major American and European, scientific journals. Such an impressive record of scholarly endeavors inspired Kefeli to write the book "Natural Growth Inhibitors and Phytohormones" which was published in Holland, in Since then, many other works were completed, including a seminal paper on "Natural Inhibitors of Phenolic and Terpenoid Nature", that appeared in 1984, in Biological Review, Cambridge, UK. Since the late eighties Dr. Maria Kalevitch (Filimonova) has been contributing to this research agenda with her investigations on growth hormones and inhibitors produced by gray molds, which infect blueberry plants. Her research substantiated the main concept of effect of these biologically active substances found in plants and other organisms as well. This theoretical framework enhanced a step further the authors' interest in directing their studies to the broader ecosystem-level and the niches played by these groups of molecules, within their respective food webs. Dr. Kalevitch's pioneering research continued through the nineties allowing her to pursue a better understanding about the role of allelopathogens produced by woody V

8 VI plants. The opportunities offered by the effect of these molecules as natural herbicides constituted the initial conceptual foundation that fostered the publication of this new book. The innovative theoretical underpinnings and ideas were presented and discussed at numerous international meetings and symposia. The theory of natural allelochemicals and their possible use as environmentally friendly herbicides was introduced also in the authors' teaching agendas, during the most recent years ( ), at different colleges and universities across the USA. Curricula in which the above mentioned principles were proposed consisted primarily in courses in, but not limited to, Environmental Science, Environmental Health, Ecology, and also student post-graduate research. This book maintains a comprehensive approach to the study of plant physiology to its end. Its educational objectives are pursued also through the presentation of research that spanned within a years time frame and didactic methods of easy implementation. The content of this work extends itself to reach many other disciplines among the environmental and agricultural sciences in particular, for its direct implications to a renovated philosophy of ecosystem management, restoration and sustainability. Its approach to the idea of plant growth stimulators-inhibitors is founded upon chemical and biological tests, as these methodologies allowed the investigators to evaluate the environmental impact of the 'regulators' under study. Appropriate laboratory techniques made ft possible to maintain the consistency necessary to validate the accuracy of the research findings. It is impossible to foresee and to predict at this time the acceptance of all the ideas and concepts presented in this work, within a broad-spectrum range of the scientific community. However, despite its limitations, this effort should be unanimously appreciated as it demonstrates a real passion to revitalize energies for the study of plant physiology and its potential to contribute to the development of a more sustainable world. Additionally, the educational perspective of the book cannot pass unnoticed as it contributes successfully to the enhancement of teaching and learning in the plant sciences. Finally, and most importantly, the work of Kefeli and Kalevitch will serve to stimulate more debates on the issue and engage the efforts of present and future researchers in an interdisciplinary manner. Professor Bruno Borsari Slippery Rock University Slippery Rock, PA USA,2002

9 VII ACKNOWLEDGMENT We would like to thank the people who shared their valuable domain knowledge and inspired us in writing this book. We express cordial thanks to the Professor M. Kh. Chailakhjan - editor of the first edition of this book. Authors express special appreciation to the members of BIOMOST, INC, USA- Mrs. Margaret Dunn and Mr. Tim Danehy for their sponsorship of our research for all these years. Our cordial thanks go to Dr. Alex E. Kalevitch without whose research, diligent and thorough preparation of this publication, the book itself will not be possible. By his heroic effort this book was born! Our book certainly will not be possible without the strong support of our colleagues from different European and American institutions whose contributions to this book always greatly and sincerely appreciated. Our thanks go to our families, especially to Mrs. Galina Mzhen, without whose patience, love, support, and total devotion this book will never come to life. Dr. Valentine Kefeli, Member of New York Academy of Sciences And Dr. Maria Kalevitch, Member of American Society of Microbiology Slippery Rock, P A, USA. 2002

10 INTRODUCTION Concept of plant growth stimulation and inhibition is tightly connected with the function of some secondary substances. They include indoles, phenols, terpenoids, adenine derivates and others. These classes of substances contain neutral and active products. The last form is present in minute amounts, and after their function is concluded in the plant, they can be easily inactivated (phytohormones and some inhibitors). Here we are discussing the products which can regulate plant growth and development, including differentiation, tissues growth and specialization, and organ interactions. Organ interactions involve such processes as: polarity, regeneration, reaction of the whole plant on the environmental factors: gravity, light, temperature, gas regim, microorganisms, and much more. Some of these substances will act as allechemicals or allelopathogens, which are secreted from the leaves and roots of the trees and herbs, and demonstrate some effect on the other plants. Allelochemicals are relative stable in the soil, and resist the neutralizing effects of microorganisms. In case of higher plants they express both: more or less resistance or sensitivity to microorganisms. In some cases microbes can change the growth activity of higher plants, examples are Fusarium that produces gibberellins which stimulate stem elongation, Taphrina - produces cytokinins and auxins which activate tumor formation or suppress apical dominance. Another great example is Botrytis, it produces abscisic acid (ABA) which is an active inhibitor in the root and shoot growth retardation. However, many aspects of hormonal biosynthesis and hormonal functions in fungus are still unclear. But now is more or less obvious that ABA regulates stomata movement; cytokinins involve in cell multiplication and greening of the leaves; auxins induce cell elongation and root regeneration; phenolic inhibitors arrest seed germination, cell elongation, and decrease the intensity of respiration. Some growth regulators synthesize in one place, and transported over the long distance, where they produce growth regulating effect. Some, like phenolic inhibitors synthesize in one certain place, and are not so movable as their precursors- amino acids or organic acids (Kefeli, Kadyrov, 1971; Kefeli, 1978; Mok and Mok, 2001; Richards et ai., 2001) Plants in the environment can be recipients and accept the effect of other plants and microorganisms. On the other hand, plants could be donors and produce growth regulating substances which could be secreted and act as botanical herbicides which inhibit selectively the growth of some species. This might be an ecological tool for the formation of natural ecosystem (Zucconi, 1996) With the help of auxins and cytokinins roots of legumes and some other species can form nodules, where the process of nitrogen fixation by Rizobium IX

11 x bacteria proceeds. At the same time bacteria of the same family, called Agrobacterium induce tumor formation and genome changes (transgenosis) via tumor formation in the other species (Lynn, Chang 1990; Chilton, 2001; Long, 2001). Thus hormones and inhibitors can be environmental factors with positive or negative signs and effects. Biosynthesis, transport and function of growth regulating substances are under the genetic and environmental control (Kende, 2001). In general these factors regulate the process of biological rhythms (so-called "Biological clock function"). Needless to say how important it is to know the targets of hormones and inhibitors action (Golden, Stayer, 2001) These targeted centers could be on the cellular level, where growth inhibitors and stimulators could interact with the cellular receptors. However, not much yet clear about how these receptors are connected with the system of effect multiplication. However this processes either integrate cellular receptor effects on the tissues targeting center or function on the organ level. The investigation of mutant plants helped to understand how genes of dwarfism block the process of biosynthesis of hormone-stimulators (like gibberellin), and activate the process of inhibitor function (ABA and some phenolic biosynthesis). Some mutants are able to produce inhibitors (ABA, ethylene) as factors of resistance to UV damage. The process of effect multiplication could be a part of the hormone-inhibitor field function, which regulates such processes as dormancy, growth, and transition to flowering, fruit production (Kefeli, 1997) No doubt not all ideas, which are presented in this book, could be accepted by researches. But we hope that even some doubtful conceptions will stimulate the plant physiologists to confirm or appose them. That was our main task when we wrote this book, to spark the research creativity and enginuity. Professor Valentine I. Kefeli Professor Maria V. Kalevitch (Slippery Rock, U.S.A.)