Biochetnistry of Fruit Ripening

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1 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.

2 First edition 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 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 ISBN (eBook) DOI / Fruit-Ripening. 1. Seymour, G. (Graham) II. Taylor, J. (J. E.) SB B '.8-dc20 III. Tucker, G. A. (Gregory A.) CIP

3 Contents Colour plate section appears between pages 406 and 407 Contributors ix 1 Introduction 1 G.A. Tucker 1.1 Respiration and energy Flavour changes Colour changes Texture changes Control of ripening Conclusions 31 References 43 2 Avocado 53 G.B. Seymour and G.A. Tucker 2.1 Introduction Physiology Biochemistry Gene expression during fruit ripening 69 References 76 3 Banana 83 G.B. Seymour 3.1 Introduction Ethylene production and respiration Carbohydrate metabolism Pigment changes Cell wall changes Phenolic compounds Organic acid and amino acid metabolism 98

4 VI Contents 3.8 Production of volatile compounds Lipids Other changes Concluding remarks 100 References Citrus fruit 107 E.A Baldwin 4.1 Commercial importance of citrus fruit Taxonomy and cultivars Harvest Postharvest General physiology Biochemical changes during development, ripening and storage Citrus biotechnology 135 References Exotics 151 J.E. Taylor 5.1 Introduction Persimmon Lychee Guava Annona species Carambola Tamarillo Passion fruit Mangosteen Feijoa Rambutan 175 References Grape 189 AK. Kanellis and K.A Roubelakis - Angelakis 6.1 Origin and distribution Grape berry morphology Fruit growth and development Growth regulation - phytohormones Postharvest physiology Handling and storage Compositional changes Phenolic compounds Aroma Cell wall metabolism 219 References 221

5 Contents vii 7 Kiwifruit 235 N.K. Given 7.1 Introduction Physiology Biochemistry Pathways and enzymes Regulation of ripening Future prospects 249 References Mango 255 C. Lizada 8.1 Introduction Fruit development and harvest maturity Ripening processes 257 References Melon 273 G.B. Seymour and W.B. McGlasson 9.1 Introduction Botany Physiological changes during development and ripening Biochemical changes during development and ripening Postharvest diseases and disorders Genetic improvement 286 References Pineapple and papaya 291 KE. Paull 10.1 The pineapple - an introduction Pineapple - physiology and biochemistry Papaya - an introduction Papaya - physiology Papaya - biochemistry 311 References Pome fruits 325 M. Knee 11.1 Introduction Physiology Biochemistry Regulation 336 References 339

6 viii Contents 12 Soft fruit 347 K. Manning 12.1 Introduction Physiology Biochemistry Metabolic pathways Hormonal regulation of development and ripening Gene expression during development and ripening 370 References Stone fruit 379 c.j. Brady 13.1 Introduction The Prunus species and their origins Peaches and nectarines (P. persica) Plum (P. domestica) Mume, or Japanese apricot (P. mume) Apricot (P. armeniaca) Almond (P. amygdalus) Tart or sour cherry (P. cerasus) Sweet cherry (P. avium) Olive (Olea europaea L) 395 References Tomato 405 G. Hobson and D. Grierson 14.1 Introduction History and classification Tomato production Fruit quality Ethylene synthesis and action Temperature stresses and theirehects on ripening The mechanism of tomato ripening Non-ripening mutants Cloning and characterization ripening-related mrnas Polygalacturonase,and its role in texture change Tomato genetic engineering Control of fruit-specific and ripening-specific gene expression 431 References 434 Glossary of botanical names 443 fud~ «5

7 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 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 Heraklion Crete MICHAEL KNEE Department of Horticulture The Ohio State University 2001 Fyffe Court Columbus Ohio USA

8 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 USA KALLIOPI A. ROUBELAKIS- ANGELAKIS Department of Biology University of Crete POBox 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

9 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

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