Biochetnistry of Fruit Ripening Edited by G.B. Seymour Horticulture Research International, West Sussex, J.E. TayIor Department of Physiology and Environmental Science, University of Nottingham, G.A. Tucker Department of Applied Biochemistry and Food Science, University of Nottingham, I~='I Springer-Science+Business Media, B.V.
First edition 1993 1993 Springer Science+Business Media Dordrecht Originally published by Chapman & Hali in 1993 Softcover reprint of the hardcover I st edition 1993 Typeset in 10/12 Palatino by EXPO Holdings, Malaysia ISBN 978-94-010-4689-3 Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the, or in accordance witj:t the terms of licences issued by the appropriate Reproduction Rights Organization outside the. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication data Biochemistry of fruit ripening/ edited by G. Seymour, J. Taylor, and G. Tucker. - Ist ed. p. cm. Includes index. ISBN 978-94-010-4689-3 ISBN978-94-011-1584-1(eBook) DOI 10.1007/978-94-011-1584-1 1. Fruit-Ripening. 1. Seymour, G. (Graham) II. Taylor, J. (J. E.) SB357.283.B56 1993 664'.8-dc20 III. Tucker, G. A. (Gregory A.) 92-38655 CIP
Contents Colour plate section appears between pages 406 and 407 Contributors ix 1 Introduction 1 G.A. Tucker 1.1 Respiration and energy 3 1.2 Flavour changes 9 1.3 Colour changes 14 1.4 Texture changes 17 1.5 Control of ripening 24 1.6 Conclusions 31 References 43 2 Avocado 53 G.B. Seymour and G.A. Tucker 2.1 Introduction 53 2.2 Physiology 54 2.3 Biochemistry 56 2.4 Gene expression during fruit ripening 69 References 76 3 Banana 83 G.B. Seymour 3.1 Introduction 83 3.2 Ethylene production and respiration 84 3.3 Carbohydrate metabolism 89 3.4 Pigment changes 90 3.5 Cell wall changes 93 3.6 Phenolic compounds 95 3.7 Organic acid and amino acid metabolism 98
VI Contents 3.8 Production of volatile compounds 98 3.9 Lipids 99 3.10 Other changes 100 3.11 Concluding remarks 100 References 101 4 Citrus fruit 107 E.A Baldwin 4.1 Commercial importance of citrus fruit 107 4.2 Taxonomy and cultivars 108 4.3 Harvest 110 4.4 Postharvest 112 4.5 General physiology 115 4.6 Biochemical changes during development, ripening and storage 120 4.7 Citrus biotechnology 135 References 137 5 Exotics 151 J.E. Taylor 5.1 Introduction 151 5.2 Persimmon 152 5.3 Lychee 159 5.4 Guava 163 5.5 Annona species 166 5.6 Carambola 169 5.7 Tamarillo 171 5.8 Passion fruit 171 5.9 Mangosteen 173 5.10 Feijoa 173 5.11 Rambutan 175 References 177 6 Grape 189 AK. Kanellis and K.A Roubelakis - Angelakis 6.1 Origin and distribution 189 6.2 Grape berry morphology 190 6.3 Fruit growth and development 193 6.4 Growth regulation - phytohormones 195 6.5 Postharvest physiology 199 6.6 Handling and storage 200 6.7 Compositional changes 201 6.8 Phenolic compounds 212 6.9 Aroma 218 6.10 Cell wall metabolism 219 References 221
Contents vii 7 Kiwifruit 235 N.K. Given 7.1 Introduction 235 7.2 Physiology 237 7.3 Biochemistry 239 7.4 Pathways and enzymes 246 7.5 Regulation of ripening 247 7.6 Future prospects 249 References 251 8 Mango 255 C. Lizada 8.1 Introduction 255 8.2 Fruit development and harvest maturity 256 8.3 Ripening processes 257 References 266 9 Melon 273 G.B. Seymour and W.B. McGlasson 9.1 Introduction 273 9.2 Botany 273 9.3 Physiological changes during development and ripening 274 9.4 Biochemical changes during development and ripening 276 9.5 Postharvest diseases and disorders 285 9.6 Genetic improvement 286 References 286 10 Pineapple and papaya 291 KE. Paull 10.1 The pineapple - an introduction 291 10.2 Pineapple - physiology and biochemistry 294 10.3 Papaya - an introduction 302 10.4 Papaya - physiology 306 10.5 Papaya - biochemistry 311 References 315 11 Pome fruits 325 M. Knee 11.1 Introduction 325 11.2 Physiology 327 11.3 Biochemistry 329 11.4 Regulation 336 References 339
viii Contents 12 Soft fruit 347 K. Manning 12.1 Introduction 347 12.2 Physiology 347 12.3 Biochemistry 354 12.4 Metabolic pathways 358 12.5 Hormonal regulation of development and ripening 364 12.6 Gene expression during development and ripening 370 References 373 13 Stone fruit 379 c.j. Brady 13.1 Introduction 379 13.2 The Prunus species and their origins 380 13.3 Peaches and nectarines (P. persica) 380 13.4 Plum (P. domestica) 390 13.5 Mume, or Japanese apricot (P. mume) 391 13.6 Apricot (P. armeniaca) 391 13.7 Almond (P. amygdalus) 392 13.8 Tart or sour cherry (P. cerasus) 392 13.9 Sweet cherry (P. avium) 393 13.10 Olive (Olea europaea L) 395 References 397 14 Tomato 405 G. Hobson and D. Grierson 14.1 Introduction 405 14.2 History and classification 406 14.3 Tomato production 407 14.4 Fruit quality 409 14.5 Ethylene synthesis and action 415 14.6 Temperature stresses and theirehects on ripening 417 14.7 The mechanism of tomato ripening 418 14.8 Non-ripening mutants 420 14.9 Cloning and characterization ripening-related mrnas 422 14.10 Polygalacturonase,and its role in texture change 425 14.11 Tomato genetic engineering 429 14.12 Control of fruit-specific and ripening-specific gene expression 431 References 434 Glossary of botanical names 443 fud~ «5
Contributors ELIZABETH A. BALDWIN United States Department of Agriculture Agricultural Research Service South Atlantic Area Citrus and Sub-tropical Products Laboratory 600 A venue Sf NW PO Box 1909 Winterhaven Florida 33881 USA COLINJ. BRADY CSIRO Division of Horticulture North Ryde New South Wales 2113 Australia DONALD GRIERSON Department of Physiology and Environmental Science University of Nottingham School of Agriculture Sutton Bonington Loughborough LE125RD THE LATE NIGEL K. GIVEN GRAEME HOBSON Horticulture Research International Worthing Road LIttlehampton West Sussex BN176LP ANGELOS K. KANELLIS Institute of Molecular Biology and Biotechnology Foundation for Research and Technology PO Box 1527 711 10 Heraklion Crete MICHAEL KNEE Department of Horticulture The Ohio State University 2001 Fyffe Court Columbus Ohio 43210 USA
x CONCEPCION C. LIZADA Post Harvest Training Research Centre University of the Philippines at Los Banos College Laguna Philippines KENNETH MANNING Horticulture Research International Worthing Road Littlehampton West Sussex BN176LP BARRY McGLASSON University of Western Sydney Hawkesbury Richmond New South Wales Australia ROBERT E. PAULL Department of Plant Molecular Physiology College of Tropical Agriculture and Human Resources University of Hawaii 3190 Maile Way Honolulu HI 96822 USA KALLIOPI A. ROUBELAKIS- ANGELAKIS Department of Biology University of Crete POBox 1470 711 10 Heraklion Crete GRAHAM B. SEYMOUR Horticulture Research International Worthing Road Littlehampton West Sussex BN176LP Contributors JANE E. TAYLOR Department of Physiology and Environmental Science University of Nottingham School of Agriculture Sutton Bonington Loughborough LE125RD GREGORY A. TUCKER Department of Applied Biochemistry and Food Science University of Nottingham School of Agriculture Sutton Bonington Loughborough LE125RD
Preface It is over 20 years since the publication of A.c. Hulme's two volume text on The Biochemistry of Fruits and thei.r Products. Whilst the bulk of the information contained in that text is still relevant it is true to say that our understanding of the biochemical and genetic mech<1nisms underlying ripening have advanced tremendously in the intervening years. For instance three of the key advances have been the elucidaton of the pathway for ethylene synthesis, the application of molecular biology to begin to unravel the genetic basis ot ripening and the manipulation of ripening by genetic engineering. This book brings the study of the biochemistry of fruit ripening up to date. The introduction is designed primarily for those readers new to the field. It provides an overview of the biochemical pathways, whose actions combine to give the multitude of physiological changes that constitute ripening. It also provides an insight into the role of ethylene in initiating and co-ordinating these various diverse biochemical pathways. The subsequent chapters deal with individual, or groups of, fruit and cover all those of current commercial importance along with some of those with future potential as economically important crops. Emphasis is given to advances in knowledge made over the last two decades, and in particular to our current understanding of the molecular biology of ripening. The book contains contributions from several experts in the field and the editors express their sincere thanks to all the contributors for their efforts and perseverance. A thorough understanding of the biochemical basis of ripening is essential if improved quality, storage and processing of fruit is to be achieved. Such improvements, by the application of genetic engineering, have already started and the scope for future advances is enormous. The prospects of producing fruit with enhanced shelf lives, better flavour and texture and higher nutritional value are good. However, our understanding of fruit biochemistry must be even further improved to assist in these aims. In particular aspects such as ethylene perception, the role of other plant growth regulators and the intricate details of the control of gene expression need to be investigated further. In providing an overview of our understanding of fruit biochemistry this book should have much to interest the plant physiologist, biochemist and molecular biologist. We hope that it will stimulate further research especially in those key areas identified by the contributors. Finally the editors would like to extend their thanks to those who helped with critical reading of parts of the final manuscript, particularly Graeme Hobson, Miguel Vendrell and Matthew Hills. G.A.T J.E.T G.BS