9780521530880cvr.qxd 15/6/07 15:06 Page 1 m u Pre-publication praise for Plant Breeding and Biotechnology: r Plant Breeding p ‘This magisterial account of the past and present of plant breeding provides a h valuable and, at times, very personal account of the development, current status y and possible futures of plant breeding. It is clear to all that we need to be able to and Biotechnology grow and distribute more food in a world which is warming, running short of K land and water and increasing in population. This book, with its extensive notes Y and bibliography, has an important part to play in the debate about the future.’ M P Professor Derek BurkeCBE (former Vice Chancellor of the University of East Anglia) l C a Societal Context and the n R Future of Agriculture V ‘…provides a comprehensive and illustrative analysis of the interface between t C science and society in the context of plant breeding, biotechnology, and the future B food situation. Professor Murphy tells a fascinating, science-based story about the y r g role of crop improvement and places the various scientific methods, including e o denis murphy genetic engineering, in proper perspective, serving as a critically important input e l o n into the debate about GMOs for food and agriculture. This book is a must-read for d h those of us who are interested in science and science policy for food and i c n e agriculture.’ t g o Per Pinstrup-AndersenH.E. Babcock Professor of Food, Nutrition and Public Bi Policy, Cornell University, and World Food Prize Laureate, 2001 a d n n a d ‘Murphy employs a fascinating blend of science, history and socio-economics to g n provide an accessible account of the development of plant breeding over the past B i d 200 years, describing the wider challenges and opportunities that lie ahead for i e o e global agriculture.’ r t B Professor Christopher J. LeaverCBE, FRS, FRSE (Sibthorpian Professor of Plant e t Science) Head oftheDepartment of Plant Science, University of Oxford c n a h Pl Cover illustration: courtesy n of the Union of Concerned : o y Scientists, USA. h l p o r u g M y Designed by Zoe Naylor PLANT BREEDING AND BIOTECHNOLOGY Societal Context and the Future of Agriculture This accessible survey of modern plant breeding traces its history from the earliest experiments at the dawn of the scientific revolution in the seventeenth century to the present dayand the existence of high-techagribusiness.Denis Murphy tells the story fromtheperspectiveofascientistworkinginthisfield,offeringarationalandevidence- based insight into its development. Crop improvement is examined from both a scientificandsocio-economicperspective,andthewaysinwhichthesefactorsinteract and impact on agricultural development are discussed. In conclusion, some concerns overthefutureofplantbreedingarehighlighted,aswellaspotentialoptionstoenable us to meet the challenges of feeding the world in the twenty-first century. This thoroughlyinterdisciplinaryandbalancedaccountwillserveasanessentialresourcefor everyone involved with plant breeding research, policy and funding, as well as those wishingtoengagewithcurrentdebatesaboutagricultureanditsfuture. Denis J. Murphy is Professor of Biotechnology at the University of Glamorgan, UK. His career in plant biotechnology research spans three decades, including ten yearsonthemanagementteamoftheJohnInnesCentre,arguablyEurope’spremier research centre in plant science. He is currently highly involved with the ongoing debate on genetically modified food and crops, both locally and internationally, providingexpertiseandadvicetonumerousorganisationsandgovernmentagencies, as well as engaging with the general public and the media. Plant Breeding and Biotechnology Societal Context and the Future of Agriculture DENIS J. MURPHY University ofGlamorgan cambridge university press Cambridge,NewYork,Melbourne,Madrid,CapeTown,Singapore,Sa˜oPaulo CambridgeUniversityPress TheEdinburghBuilding,CambridgeCB28RU,UK PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork www.cambridge.org Informationonthistitle:www.cambridge.org/9780521823890 ªD.J.Murphy2007 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithout thewrittenpermissionofCambridgeUniversityPress. Firstpublished2007 PrintedintheUnitedKingdomattheUniversityPress,Cambridge AcataloguerecordforthispublicationisavailablefromtheBritishLibrary ISBN 978-0-521-82389-0hardback ISBN 978-0-521-53088-0paperback CambridgeUniversityPresshasnoresponsibilityforthepersistenceor accuracyofURLsforexternalorthird-partyinternetwebsitesreferredto inthispublication,anddoesnotguaranteethatanycontentonsuch websitesis,orwillremain,accurateorappropriate. In the frozen midwinter of 1941–1942, the great metropolis of Leningrad began its epic 900-day siege by the encircling German army. Throughout the ruined city, thousands of people were dying of cold, starvation, and shellfire. Intheworld’soldestseedbank,attheInstituteofPlantIndustry,adedicated team of breeders and curators sought to guard and preserve their priceless samplesforposterity.Thiscollectionofover160000plantvarietieshadbeen set up in the 1920s by Nikolai Vavilov, the doyen of twentieth century plant breeding. Hardly any food reached the biologists as they maintained their protective vigil. One by one, they succumbed to starvation, surrounded by bags of edible seeds and tubers. The oats curator L.M. Rodina died, as did rice curator D.S. Ivanov, and peanut curator A.G. Shtchukin, and seven more of their heroic colleagues, one of whom even expired at his desk, working until the end. When the city was eventually liberated in January 1944, the entire collection was intact. It has since been used to supply new edible plant varieties to millions of people around the world. This book is dedicated to all the many heroes of plant breeding, both past and present, including: Norman Borlaug, Robert Carsky, Charles Darwin, ThomasFairchild,JackHarlan,MonkombuSwaminathan,NikolaiVavilov, those brave workers from Leningrad and from other more recently threatened seed banks in Asia and Africa; and, of course, the untold generations of anonymous farmer-breeders, most of whom were women. It is to you that we truly owe our daily bread. Contents Preface page xv Acknowledgements xvii Using this book xviii Nomenclature and terminology xix Abbreviations and glossary xx Introduction 1 Part I The science of plant breeding 7 1 Origins of plant breeding 9 Introduction – the development of agriculture 9 Non-intentional selection 10 Variation and selection in breeding 12 Figure 1 15 Pre-scientific empirical breeding 16 Scientific breeding 17 Beginnings of practical scientific breeding 17 Hybrids 17 Mutations 19 Quantitative genetics 21 2 Creating new genetic variation 23 Introduction 23 Hybrid crops 24 vii viii Contents Maize and other intraspecific hybrids 24 Intergenic hybrids – triticale, a new manmade crop species 28 Induced mutagenesis 29 Radiation and chemical mutagenesis 29 Somaclonal mutagenesis 31 Mutagenesis – an acceptable technology for genetic manipulation? 32 Wide crossing 33 Wide crosses in rice 34 Wide crosses in brassicas 35 3 Modern high-tech breeding 38 Introduction 38 Tissue culture technologies 39 Chromosome doubling 39 Mass propagation 41 Somatic hybridisation 42 Haploids and doubled haploids 44 Sterile plant varieties 45 Transgenesis 46 Comparison with other technologies for variation enhancement 47 Screening and selection 49 Selecting non-visible traits 49 DNA marker-assisted selection (MAS) 51 New technologies for high-tech breeding 53 Part II The societal context of plant breeding 57 4 Rise of the public sector and the US pioneers 59 Introduction 59 New technologies, old problems 60 Agricultural research as a public good 62 Emergence of public sector research in the USA 63 Morrill and the land grant institutions 65 The USDA and its botanisers 68 Extension services 71 Contents ix 5 Public sector breeding in the UK 73 Introduction 73 The UK – a laissez-faire approach 74 Rothamsted 76 John Innes Centre 77 Welsh Plant Breeding Station 78 Cambridge Plant Breeding Institute 79 Order versus chaos or control versus initiative? 81 6 Breeding goes global: the Green Revolution and beyond 83 Introduction 83 US philanthropy exported 84 CIMMYT and wheat in Mexico 86 IRRI and rice in the Philippines 87 National research organisations 88 The Green Revolution 89 Wheat 90 Rice 93 Global impact 94 The emergence of CGIAR in the 1970s 95 Part III Turmoil and transition: the legacy of the 1980s 99 7 Resurgence of the private sector 101 Introduction 101 A phoenix reborn 103 Favourable regulatory environments 105 Patents and breeders’ rights 105 The problem of broad claims 110 Startups and multinationals 112 8 Emergence of a new crop improvement paradigm 115 Introduction 115 Obsolescence and impoverishment 118 Effects on breeding programmes 121 Genetic erosion 122