World Crop Series. Available. Forthcoming titles

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1 THE POTATO CROP

2 World Crop Series Available The Grass Crop The physiological basis of production M.B. Jones and A. Lazenby The Tomato Crop A scientific basis for improvement J.G. Atherton and J. Rudich Wheat Breeding Its scientific basis F.G.H. Lupton Forthcoming titles Sugar Beet D.A. Cooke and R.K. Scott Bananas and Plantains S. Gowen Oats R.W. Welch The Groundnut Crop J. Smartt

3 THEPOTATO CROP The scientific basis for improvement Edited by Paul M. Harris Department of Agriculture University of Reading fiji Springer-Science+Business Media, B.V.

4 First edition 1978 Second edition Springer Science+Business Media Dordrecht Originally published by Chapman & HalI 1992 Softcover reprint of the hardcover 2nd edition 1992 Chapter B.D. Witney and D.C. McRae Typeset in 10112pt Times by J&L Composition Ltd, Filey, North Yorkshire Suffolk ISBN ISBN (ebook) DOI / Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK 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 UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. 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 orliability 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 The Potato crop: the scientific basis for improvementledited by Paul M. Harris. p. cm. Includes bibliographical reference.s and index. ISBN Potatoes. I. Harris, Pald M., SB211.P8P '.21--dc CIP

5 Contents Contributors Preface to the second edition Preface to the first edition Foreword xi xv xvi xviii 1 History of the potato 1 J.G. Hawkes 1.1 Introduction The status of the potato in South America Evidence for domestication - archaeological data The potato at the time of the Spanish conquest Historical and linguistic evidence The introduction of the potato into Europe The nature and exact source of the early European potato The spread of the potato into other parts of the world 9 References 11 2 Biosyst,ematics of the potato 13 J.G. Hawkes 2.1 Introduction Brief outline of potato classification Distribution and ecology of potatoes Species concepts in potatoes Crossability, sterility and breeding behaviour Endosperm balance numbers Cytology of potato species Chemotaxonomic relationships of potatoes Evolutionary divergence and genome differentiation in wild potato species Potato evolution Evolutionary relationships of cultivated potato species 57 References 60

6 vi Contents 3 Structure and development of the potato plant 65 Elizabeth G. Cutter 3.1 Introduction Development of the seedling The root system The leafy shoot The stolon The tuber Dormancy and its control Development and structure of the flower Embryogenesis Tissue culture Genetic engineering Conclusions 145 Acknowledgements 146 References Mineral nutrition 162 P.M. Harris 4.1 Introduction Nutrient accumulation and use Relations between mineral nutrients, growth and yield Fertilizer requirements Some factors modifying the yield response to fertilizers Fertilizer use and potato quality Conclusions 207 References Water relations and growth of potatoes 214 P.J. Gregory and L. P. Simmonds 5.1 Introduction Growth and water use Y.ield and water supply Root growth The physiological basis for the drought-sensitivity of potato growth Water supply and tuber quality 236 Acknowledgements 241 References Seed tuber production and management 247 E.J. Allen, P.l. O'Brien and D. Firman 6.1 Introduction Growth and development of the seed tuber Seed multiplication and certification Seed production and utilization 278

7 Contents vii 6.5 True seed Acknowledgements References Plant density 292 E.l. Allen and D.C.E. Wurr 7.1 Introduction Units of density Spatial arrangement Effects of plant density 312 References Breeding new varieties 334 P.D.S. Caligari 8.1 Introduction Breeding methods Breeding objectives The potential role of other breeding techniques 362 References Weed control 373 P.l. W. Lutman 9.1 Weeds in potatoes Weed biology Weed control Recommendations for the use of herbicides Potato haulm destruction Potatoes as weeds 396 References Disease aspects of potato production 403 G.A. Hide and D.H. Lapwood 10.1 Introduction Effects of diseases Survival and spread of pathogens Control of diseases Future prospects 431 References Pest aspects of potato production Part 1 The nematode pests of potatoes 438 K. Evans and D.L. Trudgill 11.1 Introduction to their biology Effects of root-feeding nematodes on growth and yield Interactions with other organisms Methods of decreasing nematode damage Future developments in nematode research 462 References 466

8 Vlll Contents Part 2 Insect pests 476 K. V. Raman and E. B. Radcliffe 11.6 Introduction Pest biology, damage and distribution Pest control measures Integrated pest management (IPM) 495 References Tuber quality 507 R. M.J. Storey and H. V. Davies 12.1 Introduction Quality associated with the fresh tuber Quality associated with internal factors Aspects affecting quality of the cooked potato Processing quality Irradiation Concluding comments 551 References Mechanization of crop production and handling operations 570 B.D. Witney and D.C. McRae 13.1 Introduction Seedbed preparation Fertilizer application Potato planting Harvesting Handling and grading Conclusions and future prospects 604 References The physics and physiology of storage 608 W. G. Burton, A. van Es and K.J. Hartmans 14.1 Introduction Physics Wound healing and curing Dormancy, sprout growth and ageing Changes in composition during storage 677 References Potato production in the tre,pics 728 D.J. Midmore 15.1 Introduction Production statistics for the tropics and sub tropics Tropical highland potato production (> 1500 m) Tropical lowland potato production «1500 m) - constraints and solutions 742

9 Contents ix 15.5 Tropical lowland potato production - two practical examples General comments and future 779 References Potato production in the context of the world and farm economy 794 D.E. Horton and 1.L. Anderson 16.1 Global patterns and trends Potatoes in the food systems of developing countries Potatoes in developed countries 806 References Principles of agronomy and their application in the potato industry 816 E.J. Allen and R. K. Scott 17.1 Crop growth and development The use of agronomic principles in understanding the effects of agronomic treatments Relevance of agronomic understanding to improvements in the potato industry 873 Acknowledgements 878 References 878 Index 882

10 Contributors E.J. Allen Cambridge University Farm Huntingdon Road Cambridge CB30LH, UK J.L. Anderson The Scottish Agricultural College Rural Resource Management Division 42 South Oswald Road Edinburgh EHO 2HH, UK Dr W.G. Burton deceased P.D.S. Caligari Department of Agricultural Botany Plant Science Laboratories University of Reading Whiteknights PO Box 221 Reading RG6 2AS, UK E.G. Cutter School of Biological Sciences Department of Cell and Structural Biology University of Manchester Williamson Building, Oxford Road Manchester M13 9PL, UK H.V. Davies Scottish Crop Research 1nstitute 1nvergowrie Dundee DD2 5DA, UK

11 XlI Contributors A. van Es Pootakkerweg BX Wageningen, The Netherlands K. Evans Crop and Environment Protection Division Entomology and Nematology Department Rothamsted Experimental Station Harpenden Herts AL5 2JQ, UK D. Firman Cambridge University Farm Huntingdon Road Cambridge CB3 OLH, UK P.J. Gregory CSIRO Dryland Crops and Soils Research Unit Private Bag PO Wembley WA6014 Western Australia P.M. Harris Department of Agriculture University of Reading Earley Gate PO Box 236 Reading RG6 2AT, UK K.J. Hartmans IBVL PO Box) Wageningen, The Netherlands J.G. Hawkes c/o School of Continuing Studies University of Birmingham PO Box 363 Birmingham BI52TT, UK G.A. Hide Crop and Environment Protection Division Plant Pathology Department Rothamsted Experimental Station Harpenden Herts AL5 2JQ, UK

12 Contributors Xlll D.E. Horton ISNAR PO Box The Hague, 2509 Al The Netherlands D.H. Lapwood Crop and Environment Protection Division Plant Pathology Department Rothamsted Experimental Station Harpenden Herts AL5 2IQ, UK P.I.W. Lutman Crop Management Department AFRC-IACR Rothamstead Experimental Station Harpenden, Herts, AL5 2IQ D.C. McRae Scottish Centre of Agricultural Engineering Bush Estate Penicuik Midlothian, EH260PH, Scotland, UK D.I. Midmore The Asian Vegetable Research and Development Center PO Box 42, Shanhua Tainan, Taiwan, Republic of China P.I. O'Brien Cambridge University Farm Huntingdon Road Cambridge CB3 OLH, UK E.B. Radcliffe Department of Entomology College of Agriculture University of Minnesota 219 Hodson Hall, 1980 Folwell Avenue St Paul, Minnesota , USA K.V. Raman Nematology and Entomology Department International Potato Center PO Box 5969 Lima, Peru

13 XIV Contributors R.K. Scott Department of Agriculture School of Agriculture Nottingham University Sutton Bonington Loughborough Leics LEl2 5RD, UK L.P. Simmonds Department of Soil Science University of Reading London Road Reading RGI 5AQ, UK R.M.J. Storey Potato Marketing Board Broadfield House 4 Between Towns Road Cowley Oxford OX43NA, UK D.L. Trudgill Scottish Crop Research Institute Invergowrie Dundee DD2 5DA, UK B.D. Witney Scottish Centre of Agricultural Engineering Bush Estate Penicuik Midlothian, EH260PH, Scotland, UK D.C.E. Wurr Horticulture Research International Wellesbourne Warwick CV35 9EF, UK

14 Preface to the second edition Research and publications on the potato crop have burgeoned since the first edition of this book was published in However, the warm reception of the first edition suggested that it had a useful part to play in promoting the scientific basis for understanding and improving the yield and quality of the crop. Since the first edition was out of print and a second reprint would not have taken into account the contributions made by research over the intervening years, it became obvious that a complete revision was necessary. There was, in particular, a need to take account of the rapid extension of interest in the crop into climates and farming systems with which it has not been traditionally associa,ted. Those involved with the crop will be sadly aware that a number of contributors to the first edition are no longer with us. Their contribution to our knowledge of the crop will however be a permanent legacy of their achievement. I would like to thank all those who have contributed to the book for their willingness to cooperate in the difficult task of bringing their particular subject up to date. This is even more noteworthy for the pressure of time appears to be almost an order of magnitude higher than it was when the first edition was tackled. Under the circumstances it is perhaps not surprising that not all the initial objectives have been achieved; that most of them have is a tribute to the contributors, to whom I would like to dedicate this edition of the book. Paul M. Harris University of Reading November 1991

15 Preface to the first edition One of the major objectives of this book on the potato has been to review the knowledge which is fundamental to successful potato production and upon which further improvement must be based. Underlying this objective is the belief that high yields, efficiently obtained are of prime importance to producers and consumers and to research workers who serve the interests of both. Despite the paradox that the greatest financial returns are often achieved when yields are lower than normal, profitability through scarcity is not in the long-term interest of either producers or consumers. This attempt to review the knowledge on which the scientific production of potatoes is based has necessitated seeking contributions from a fairly large number of specialists, which is a well-recognized method of dealing with subjects which are too large to be considered in the necessary depth by anyone author. However this tendency to specialize within fairly narrow limits carries with it the attendant dangers that few research workers see the complete picture; on the other hand the producer is concerned with the whole system of producing a crop but may not have sufficient knowledge of particular components of the system which may s i g n i f iaffect c a n the ~ l y efficiency with which his enterprise is operated. It is the hope that this book, read in its entirety, will provide the basic biological principles on which more productive and economic systems of potato production can be based. Inevitably the views presented in this book will have been coloured by the particular experiences and enthusiasm of the contributors, and no doubt much has been omitted that might have been included and possibly vice versa; however the overall balance is the responsibility of the editor and any shortcomings in this respect should be attributed to him. It is my hope that anyone reading the book who h,as constructive suggestions to make will have no hesitation in contacting the editor or contributors. I am indebted to Professor Rober,ts for asking me to edit this book and to the contributors who have so willingly cooperated in its production. Thanks are also due to my wife and family for their patient forebearance

16 Preface to first edition xvii during the book's preparation. Finally I would like to dedicate this book to all those who have contributed to our knowledge of this fascinating, useful and enjoyable vegetable. University of Reading June, 1977 P.M.H.

17 Foreword The first edition of this book was published twelve years ago. It established the first of a continuing series of monographs on important crops. This series arose from the perception that, while the disciplines which contribute to improving crop production are becoming more specialized, so it is becoming increasingly necessary to integrate this knowledge in order to improve production systems. Farmers, growers and agronomists have always had to try and do this; but it is also important for the scientific specialists who serve them to have some grasp of cognate disciplines, so that an understanding of the wider context can help them to target their contribution more effectively. It is no longer really possible for one author to write an up-to-date book which adequately and critically deals with all facets of a crop. An account which satisfactorily covers the field requires the efforts of a number of specialists. And if this account is to be coherent and well-balanced it needs careful planning and editing. The first edition of this book was based on this simple principle and it became the model for the series. The reception and demand for the first edition suggests that many people concerned with the potato crop found this approach helpful. In the meanwhile, considerable progress has been made in some areas; some of the original authors have moved on to other interests, and sadly, others have died. A new edition was clearly called for and the opportunity has been taken for a complete revision. The underlying philosophy which concentrated on unde.rlying principles, which consequently are not limited by geographical boundaries, remains the same. But the contributions of several new authors, and the recognition of an increasing interest in the crop in the tropics, has given this new edition an even greater international flavour. I believe that anyone who is seriously interested in this important crop will wish to have a copy near to hand. Eric H. Roberts World Crop Series Editor November 1991