THE ROLE OF ASCORBIC ACID IN GROWTH, DIFFERENTIATION AND METABOLISM OF PLANTS ADVANCES IN AGRICULTURAL BIOTECHNOLOGY Related titles previously published Akazawa T., et aI., eds: The New Frontiers in Plant Biochemistry. 1983. ISBN 90-247-2829-0 Gottschalk W. and Muller H.P., eds: Seed Proteins: Biochemistry, Genetics, Nu tritive Value. 1983. ISBN 90-247-2789-8 Marcelle R., Clijsters H. and Van Poucke M., eds: Effects of Stress on Photosynthesis. 1983. ISBN 90-247-2799-5 Veeger C. and Newton W.E., eds: Advances in Nitrogen Fixation Research 1984. ISBN 90-247-2906-8 The Role of Ascorbic Acid in Growth, Differentiation and Metabolism of Plants by the late J.J. CHINOY Gujarat University, India edited by N.J. CHINOY Gujarat University, India co-edited by I.e. DAVE Bhabha Atomic Research Centre Bombay, India Y.D. SINGH Saurashtra University Gujarat, India O.P. SAXENA Gujarat University, India A.V. VYAS M.G. Science Institute Ahmedabad (Gujarat), India 1984 MARTINUS NIJHOFF lOR W. JUNK PUBLISHERS a member of the KLUWER ACADEMIC PUBLISHERS GROUP THE HAGUE / BOSTON / LANCASTER Distributors for the United States and Canada: Kluwer Boston, Inc., 190 Old Derby Street, Hingham, MA 02043, USA for all other countries: Kluwer Academic Publishers Group, Distribution Center, P .O.Box 322, 3300 AH Dordrecht, The Netherlands Library of Congress Cataloging in Publication Data Library of Congress Cataloging In Publication Data Chinoy, J. J., d. 1978. The role of ascorbic acid in growth, differentiation, and metabolism of plants. (Advances in agricultural biotechnology) Includes index. 1. Plants, Effect of Vitamin C on. I. Chinoy, N. J. II. Title. III. Series. QK753.V58c48 1984 581.19'26 83-25104 ISBN-13: 978-94-010-3715-0 e-ISBN-13: 978-94-010-3713-6 001: 10.1007/978-94-010-3713-6 Copyright © 1984 by Martinus NijhofflDr W. Junk Publishers, The Hague. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publishers, Martinus NijhofflDr W. Junk Publishers, P.O. Box 566, 2501 CN The Hague, The Netherlands. v CONTENTS Preface & Acknowledgements vii I. ASCORBIC ACID: CHEMISTRY, PROPERTIES, BIOSYNTHESIS AND ASSAY 1. Structure and properties of free 1 and bound Ascorbic Acid 2. Biogenesis, detection and deter- 28 mination of free and bound Ascorbic Aeid II. ASCORBIC ACID IN PLANT GROWTH AND DEVELOPMENT 3. Photosynthesis and Respiration 68 4. Germination, Juvenile and Vegetative 100 growth 5. Reproductive Growth 150 6. Senescence 189 7. Root Nodules 195 III. ASCORBIC ACID AND STRESS PHYSIOLOGY 8. Drought resistanee and presowing 212 treatments 9. Radiation 265 IV. ASCORBIC ACID AND METABOLIC REGULATION 10. Polyphenols 277 11. Enzyme aetion 283 12. Regulation 303 Subject Index 320 VII PREFACE There is a paucity of information on the dynamics of Ascorbic Acid (AA) turnover in relation to germination, metabolism, growth, differentiation and development of a plant and in those undergoing stress of various types. in presowing treatment of seeds etc. The turnover of AA plays an important role during the juvenile phase of growth of a plant and has a significant bearing on its subsequent growth, development and maturation. The beneficial effect of presowing treatment of seed with Ascorbic Acid (AA) + H2 O2 highlights the validity of the AA-nucleic acid protein metabolism concept of growth and development of plan ts. During the course of the last 30 years, work has been undertaken by the author and his collaborators on the meta bolic drifts of regulatory substances during juvenile, vegetative, reproductive and senescent phases. The most important of these growth regulatory substances was found to be Ascorbic Acid. The dynamiC role of AA turnover is revealed by its control of rates of metabolic processes as well as those of enzymic reactions which paves the way to "New Genetics". A reference to the role of AA in metabolism and growth of the animal cell has been specially made with a view to emphasize the oneness of the fundamental reactions in both plants and animals. Fundamental processes viz. the onset of reproductive phase, microsporogenesis. megasporogenesis, embryogenesis, and others in the plant as well as spermato genesis, embryogenesis, muscle metaholism and those of a number of other tissues in the animal reveal identical metabolic machinery powered by the electron energy of the free radicals formed by the falling apart of the complexing partners of the charge transfer complex hetween AA and macro molecules. Thus at the molecular and suhmolecular levels, the plant and the animal systems are very similar in their ascorbate turnover pattern. VIII The foresoins _onosraph on - 'THE ROLE OF ASCORBIC ACID IN GROWTH, DJ:'FFERENTIATION AND METABOLISM OF PLANTS' is. therefore, a special treatise on a theme which has not heen hitherto fully exploited in detail. Morphological, bio chemical. biopbysical and histochemical evidences have been presented in this compilation to elucidate the electron donating role of AA in many a plant process. To the best of our knowledge. this book should rank as one amongst the few availab~e monographs elucidating the role of Ascorbic Acid in various facets of plant life. Moreover, the work incorporated in the book has considerable practical importance for obtaining higher yield of crops, better growth of plants and for producing those which are able to withstand stress conditions. Ahmedabad, 1983 Editor Co-Editors IX ACKNOWLEDGEMENTS I place on record .y sincere gratitude to the University Graftts Commission, New Delhi, India, for allowing me to compile this monograph and for offering all necessary faci lities for carrying out extensive research on multifarious roles of AscorbiC Acid in plant growth and productivity. My grateful thanks are due to Professors O.P. Garg of Depart.ent of Botany. Punjab University, Chandigarh, India. and M.M. Laloraya of Department of Life Sciences, Indore University. Indore (MP). India. for providing timely material for compilation of Chapters 7 and 10 respeetively. I deem it a pleasure to express my appreciation to Professors F.A. Loewus, M. Miehniewicz,. D.I. Arnon, L.W. Mapson, A. Szent Gyorgyi. B. Aberg. K.K. Nanda. G.S. Sirohi. K. Mothes. M. Reid. M. Kutacek. O. Stocker, I. Yamaz~ki. P. Schopfer. S. Tonzig and many others for making available their reprints. My special thanks go to Mr. R.K. Modi for painstakingly typing out the monograph; Mr. N.A. Shah for the expert handling of photographic prints; the artists Mr. Somabhai Dholakia and Mr. N.V. Shah for their untiring efforts; and all the staff of the Department of Botany and The Gujarat University for helping me in various ways during the compilation of this monograph. Finally. I would like to express profound thanks and appreciation to my dear wife, Roda for her dedication, silent forbearance, help and encouragement without which this work would have remained incomplete. Professor J.J. Chinoy Professor of Botany and Director University School of Sciences Gujarat University Ahmedabad - 380 009 Ahmedabad. 1978 x The Editor and Co-editors of this _ana graph acknowledge with sincere gratitude the help rendered by all research students of Professor 3.J. Chinoy, especially Dr. J.N. Patel, Dr. P.V. Vaishnav, Dr. K.C. Bhatt, Dr. D.B. Jadeja, Dr. N.S. Sharma and Dr. S. Verghese after his sad and sudden demise on 12th May, 1978, in the ardous task of proof reading and checking of references. The willing assistance of Dr. Kersi Fanibunda of Newcastle-Upon-Tyne, UK; Dr. Lovji D. Cama of New Jersey, USA; Dr. M.V. Rao and Dr. R.J. Verma, Department of Zoology, Gujarat University, in proof reading is highly appreciated. The timely help rendered in typing out the manuscript by Mr. V.T. Viswanathan is greatly appreciated. The Editor and Co-editors express grateful thanks to Madam Roda ~. Chinoy for per_itting the free and full access to all the relevant papers, data and literature of Late Professor J.J. Chinoy to enable uS to Iinalize the monograph. We also hasten to mention that the reference to work on Ascorbic Acid Metabolism in Animals is the research work carried out by Dr. (Miss) N.J. Chinoy. We have made extensive use of Dr. O.P. Saxena's review paper on 'Presowing Treatment of Seeds' while compiling Chapter B for which we owe grateful thanks to him. And finally, we would like to thank one and all in our respective families and from the Departments of Botany and Zoology, Gujarat University, Ahmedabad, India, for helping us in various ways. Editor Dr. (Miss) N. J. Chinoy Gujarat University, India Co-Editors Dr. I. C. Dave Dr. O. P. Saxena Bhabha Atomic Research Centre Gujarat University, India Bombay, India. Dr. Y. D. Singh Dr. (Miss) A. V. Vyas Saurashtra University M.G. Science Institute Gujarat, India. Ahmedabad, Gujarat, India XI ABBREVIATIONS Ascorbic acid Free Radical of ascorbic acid Ascorbic acid free radical AA - MM ~ Ascorbic acid - macromolecule complex AAO • Ascorbic acid oxidase AAU • Ascorbic acid utilization AgN03 Silver nitrate ASG Ascorbic acid bound or ascorbigen ATP Adenosine triphosphate Bz Benzimydazole C Control (non-vernalized) Ce .. Cerium * Chlorophyll Light excited chlorophyll Co .. Cobalt Col Colchicine Cs Cesium CTC Charge transfer complex Cu Copper cv Cultivar DCMU -3-(3'-4' -dichlorophenyl)-l, I-dime - thylurea DCPIP 2, 6-Dichlorophenol indophenol DCPIPH2 2, 6 Dichlorophenol indophenol reduced DDC Diethyldithiocarbamate DHA Dehydroascorbic acid DKG 2, 3 - diketogulonic acid DNPH 2, 4 - Dinitrophenylhydrazine DPPH' .. a, a - diphenyl - B - picryl hydrazyl DW Distilled water e. value .. Extinction value E,£ Enzyme, Energon EI Redox potential o EDTA Ethylene diamine tetra acetic acid EMH Enzyme mobilizing hormone .. EPR Electron paramagnetic resonance
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