Ebook Plant breeding and biotechnology societal context and the future of agriculture part 1

Ebook Plant breeding and biotechnology societal context and the future of agriculture part 1. This page intentionally left blank PLANT BREEDING AND BIOTECHNOLOGY Societal Context and the Future of Agriculture This accessible survey of modern plant breedin.Ebook Plant breeding and biotechnology societal context and the future of agriculture part 1

Societal Context and the Future of Agriculture

This accessible survey of modern plant breeding traces its history from the earliestexperiments at the dawn of the scientific revolution in the seventeenth century to thepresent day and the existence of high-tech agribusiness Denis Murphy tells the storyfrom the perspective of a scientist working in this field, offering a rational and evidence-based insight into its development Crop improvement is examined from both ascientific and socio-economic perspective, and the ways in which these factors interactand impact on agricultural development are discussed In conclusion, some concernsover the future of plant breeding are highlighted, as well as potential options to enable

us to meet the challenges of feeding the world in the twenty-first century Thisthoroughly interdisciplinary and balanced account will serve as an essential resource foreveryone involved with plant breeding research, policy and funding, as well as thosewishing to engage with current debates about agriculture and its future

De n i s J Mu r p h y is Professor of Biotechnology at the University of Glamorgan,

UK His career in plant biotechnology research spans three decades, including tenyears on the management team of the John Innes Centre, arguably Europe’s premierresearch centre in plant science He is currently highly involved with the ongoingdebate on genetically modified food and crops, both locally and internationally,providing expertise and advice to numerous organisations and government agencies,

as well as engaging with the general public and the media

Plant Breeding and Biotechnology

Societal Context and the Future

of Agriculture

DENIS J MURPHYUniversity of Glamorgan

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press

The Edinburgh Building, Cambridge CB2 8RU, UK

First published in print format

ISBN-13 978-0-521-82389-0

ISBN-13 978-0-511-34280-6

© D J Murphy 2007

2007

Information on this title: www.cambridge.org/9780521823890

This publication is in copyright Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press.

ISBN-10 0-511-34280-2

ISBN-10 0-521-82389-7

Cambridge University Press has no responsibility for the persistence or accuracy of urls for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Published in the United States of America by Cambridge University Press, New York

www.cambridge.org

hardback

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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.

In the world’s oldest seed bank, at the Institute of Plant Industry, a dedicated team of breeders and curators sought to guard and preserve their priceless samples for posterity This collection of over 160 000 plant varieties had been 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, Thomas Fairchild, Jack Harlan, Monkombu Swaminathan, Nikolai Vavilov, 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.

Preface page xv

Introduction – the development of agriculture 9

Maize and other intraspecific hybrids 24

Intergenic hybrids – triticale, a new manmade crop species 28

Comparison with other technologies for variation enhancement 47

Part II The societal context of plant breeding 57

4 Rise of the public sector and the US pioneers 59

Emergence of public sector research in the USA 63

5 Public sector breeding in the UK 73

Order versus chaos or control versus initiative? 81

6 Breeding goes global: the Green Revolution and beyond 83

Part III Turmoil and transition: the legacy of the 1980s 99

8 Emergence of a new crop improvement paradigm 115

Adapting crops to commercial agronomy 122

A technology focus based on short-term profitability rather

Privatisation, integration and globalisation 127

The Agricultural Development and Advisory Service 133

Dangerous liaisons – partnerships with the private sector 146

Gene transfer within and between plant genomes 164

Genetic manipulation or manipulation of genomes? 165

Agbiotech today – the worst of all possible worlds? 168

Rise and fall of the ‘life-sciences’ biotech business model 172

12 The future of transgenic crops I: improving

Management, segregation and other challenges 186

13 The future of transgenic crops II: improving

Part V Increasing global crop production:

14 Feeding the world – fallacies and realities 213

Population, economic growth and food production 215

Reclaiming abandoned and set-aside land 221

15 The roles of management, subsidies and breeding in crop

Agricultural overproduction and subsidies 233

Subsidies and tariffs stifle development 233

Part VI Plant breeding in the twenty-first century 239

16 The future of international plant breeding 241

Vulnerability of international seed banks 254

UC Davis and the Iranian National Seed Bank 254

ICARDA and the Iraqi National Seed Bank 255

Rebuilding agriculture across the world 257

International patrimony or restricted property? 259

17 Rebalancing our approach to crop improvement 264

Rebalancing the public debate on agriculture 269

Empowering and recruiting the private sector 277

A new market-based public sector paradigm 281

Open access technologies in plant breeding 283

Domesticating new crops – an alternative to transgenesis 288

Innovative applied R&D – the USA leads (again) 290

This book is an evidence-based and, in places, a personal account of the development

of scientific plant breeding over the past two hundred years The work is informed by

my background and experiences as a biologist who, while largely trained in the UK,has also worked extensively in the USA, Germany, Australia and, more recently, inthe Far East It is the story of how breeding evolved from an empirical endeavour,practised for millennia by farmers and amateur enthusiasts, to become the globalisedcorporate agribusiness enterprise of today I was moved to write this account afterspending over two decades working at the interface of academic plant science and itspractical application in crop breeding During this time I have witnessed the steadyerosion of plant breeding as a worthwhile and respected aspect of plant science,especially in the public sector One of my principal motives in writing the book is toraise the profile of plant breeding as a valued and useful profession I also wish tohighlight some of the many imbalances that now bedevil our approach to breeding,some of which have coloured today’s often contentious discourse on agriculture andcrop improvement in general

There are many misapprehensions, among scientists and the general public alike,about the way that plant breeders go about their business In particular, thesupposedly revolutionary nature of the ‘new’ (actually now more than two decadesold) technologies of genetic engineering has been exaggerated by virtually everybodyinvolved in the debate, whether they be researchers, politicians, agbiotech companies

or anti-GM (genetic manipulation) campaigners The current fixation on this at timesoverhyped phenomenon is coupled with a worrying dearth of knowledge andunderstanding about the many other (non-transgenic) forms of plant breeding which,

as I will show, can in principle be subjected to many of the same objections that arelevelled against GM technologies So, why is it that this particular aspect of plantbreeding is deemed so threatening that it can elicit violence and disorder among oftenidealistic and well-intentioned anti-GM activists, while the same people know little ofthe rudiments of plant breeding in the wider sense? And it is not just anti-GMcampaigners who have little knowledge of the broader socio-scientific dimensions of

xv

plant breeding; a similar charge can be levelled against many people in the broaderrealms of science, politics, the media etc.

In Part I, we will begin to address these issues by discussing the basic scientificbackground of plant breeding These three chapters are the only ones that focus onscience per se The remaining five parts of the book are devoted to examining theinterface between science/technology and society, and the manner in which theseforces have mutually influenced each other in the case of plant breeding and theproduction of improved crops In this analysis, I will take it as read that science andtechnology are deeply embedded in the wider socio-economic milieux from which theyboth arise This is particularly true in the case of plant science and its applications viavarious technologies to effect improvements in crop performance As we will see, an

‘improvement’ is normally so defined by the improver; hence one person’simprovement might even be another’s curse Science and society are respectivelymade up of many players, all of whom interact with and affect one another, often insubtle ways that are not always obvious to the casual observer In the case of GMcrops, the peculiar, and unusually contentious, trajectory of this technology has beendetermined by interactions between a host of factors including scientific discoveries(how to transfer genes), legislative measures (patenting plants), the economicenvironment (privatisation), political opportunism (policy based on pressure groups),ideology (policy based on belief systems), and so on I will show how it was aparticular conjunction of circumstances in the 1980s and 1990s that has led to thecurrent, and arguably inappropriate, domination of the agbiotech/GM phenomenon

in both scientific and public discourse A key message that I wish to convey is the need

to rediscover a sense of perspective in our attitude to crop improvement and to raiseour gaze beyond the narrow confines of the GM debate, so that we can behold the realchallenges and opportunities that confront international agriculture in the twenty-firstcentury

I am indebted to the many friends and colleagues who have, wittingly or not,inspired and assisted me in various ways during the writing of this book Thisincludes those colleagues at the John Innes Centre and University of Glamorganwith whom I have had numerous fruitful discussions over the past 15 years Specialthanks are due to Eddie Arthur and Ray Matthias with whom I tried (and failed) tointerest funding agencies in the domestication of new crops – rest assured, our timewill come As an initially somewhat reductionist molecular biologist, I waschallenged in the early 1990s by Colin Law who remained a true believer intraditional plant breeding, and a sceptic of the many chiliastic claims of agbiotech Iguess some of his sentiments must have eventually rubbed off on me, as this bookshows Other colleagues, including Ian Bartle, Gerry Roberts and Colette Murphyprovided valuable feedback on various drafts of the manuscript, as did variousanonymous referees Finally, many thanks to Katrina Halliday and the staff atCambridge University Press for their patience and encouragement during thegestation of this project.

Denis J Murphy

Glamorgan, Wales, April 2006

xvii

This book is aimed at several audiences, from botanists to economists, and frombusiness people to agronomists Each group of readers will have different technicalbackgrounds and different types of expertise The book is therefore written on threelevels, namely the main text, a series of more than 900 endnotes, and a bibliography.For the general reader or specialist in other areas, the main text of each chaptershould suffice to convey its key message However, for those wishing to follow uppoints in more detail, the endnotes provide additional information that is in turnlinked to a comprehensive bibliography of over 750 citations, mostly from the peer-reviewed primary literature Wherever possible, I have also provided web links to themany articles that are now available online Many of the better scientific journalsnow make their articles freely available on the Internet within a year of publication,and such web links tend to be relatively stable These primary research articles canoften be surprisingly accessible, even to the interested layperson, and I recommendinterested readers to consult at least one or two examples.

Secondary literature sources, e.g scholarly reviews, government reports,conference papers etc., are also often available on the Internet and can frequently

be useful resources, especially for the technical specialist from a different field Sucharticles generally give a broader perspective than primary research papers, but maynot necessarily be peer reviewed ‘Tertiary’ sources, including newspaper andmagazine articles, are rarely peer reviewed Such articles tend to be more ‘fresh’ andaccessible in their content, but can also be less factually reliable, and rarely provide abroad overview of the topic in question Tertiary sources also tend to be moreephemeral in their Internet locations (hence, caveat lector and my apologies inadvance if some of the web links no longer work) Nevertheless, newspaper andmagazine articles often add a welcome degree of colour and immediacy to adiscussion that contrasts favourably with the more sober and restrained tone of mostmainstream scientific literature

xviii

Measurements and dates

The metric system is used throughout for all physical measurements All prices aregiven in US dollars ($) and, unless stated otherwise, they relate to the period inquestion, i.e prices are not corrected to present-day values Some historical dates areexpressed as BCE (before common era) or CE (common era) All unqualified datesrefer to CE

Initials and acronyms

I have tried to forebear, as much as possible, from using unfamiliar initials andacronyms in the main text Where this is impractical, I give the full version of eachterm in the text when it is first used A full list of such terms, plus some additionalexplanation of their significance, is also given in the Abbreviations and glossarysection (see below)

Industrial and developing countries

In describing the major global economic blocs, the term ‘industrial’ is used todescribe those countries that have already completed a thorough industrialisingprocess In many cases such countries are now at a post-industrial stage ofdevelopment Included here are the major globalised economies of North America,Europe, Australasia and Japan I use the term ‘developing’ to describe thosecountries in which industrialisation is still proceeding, albeit often at an advancedstage This group includes the Asian giants India and China, as well as most of theremainder of Asia, Africa and Latin America This terminology is by no meansperfect and there will always be exceptions Neither is it meant as a value judgementsince all countries are always in the process of some sort of development But itremains, withal, a useful shorthand

xix

ADAS Agricultural Development and Advisory Service (UK) – The former

public sector agricultural advisory service for England and Wales.ADAS was gradually privatised during the 1990s until it became aprivate company in 1997

the UK government strategic advisory body on biotechnologyissues affecting agriculture and the environment Established in

2000, the AEBC was wound up in 2005, following criticisms of itsnarrow remit and dissension among members

BBSRC Biotechnology and Biological Sciences Research Council (UK) –

the major UK public sector funding agency for research inbiological sciences, with an annual budget of about $550 million

CAMBIA to ‘foster democratic innovation in applications ofbiological technologies to sustainable development’

Bt Bacillus thuringiensis – a bacterium that produces a variety of

insecticidal protein toxins Bt sprays (containing bacterial sporesand toxin crystals) are regularly used as insecticides by organicfarmers, while the Bt toxin gene has been added to some crops toprovide inbuilt insect protection

CAMBIA Center for the Application of Molecular Biology to International

Agriculture – non-profit, Australian-based scientific organisationworking for the development of new open access technologies forcrop improvement across the world

xx

CGIAR Consultative Group on International Agricultural Research – an

alliance of countries, international and regional organisations, andprivate foundations that supports 15 Research Centres TheCentres work with national agricultural research systems and civilsociety organisations including the private sector and generateglobal public goods that are freely available to all CGIAR

ICARDA, ICRAF, ICRISAT, IFPRI, IITA, ILRI, IPGRI, IRRI,IWMU, WARDA and WFC

CIAT Centro Internacional de Agricultura Tropical (Columbia) – one of

the CGIAR crop improvement centres

CIFOR Center for International Forestry Research (Indonesia) – one of

the CGIAR crop improvement centres

CIMMYT Centro Internacional de Mejoramiento de Maiz y Trigo

(Inter-national Maize and Wheat Improvement Center Mexico) – one ofthe CGIAR crop improvement centres

CIP Centro Internacional de la Papa (Peru) – one of the CGIAR crop

improvement centres

CSIRO Commonwealth Scientific and Industrial Research Organisation –

the principal public sector research organisation in Australia thatcovers agribusiness; information, manufacturing and minerals; andsustainable energy and environment CSIRO manages severalresearch centres that work on crop-related topics One of the bestknown of these is CSIRO Plant Industry in Canberra, which isespecially noted for its research on plant molecular and develop-mental biology The annual budget is about $700 million

Cultivar a cultivated variety of a crop – such varieties have normally been

selected by breeding and are adapted for a particular agriculturaluse or climatic region

formerly known as MAFF, DEFRA was created in 2001 in thewake of the BSE scandal but lost its role in food safety to the newFood Standards Agency (FSA)

EMBRAPA Empresa Brasileira de Pesquisa Agropecua´ria–Brazilian

Agricul-tural Research Corporation

EST expressed sequence tag – a small portion of a gene that can be used

to help identify unknown genes and to map their positions within agenome

Ex situ

conservation

the maintenance of biological specimens away from their normalhabitat, normally under closely controlled conditions, such as inarboretums (trees), and botanical (plants) or zoological (animals)gardens The term also refers to the keeping of stocks, such asseeds, cuttings, or other propagules in germplasm repositories.FAO Food and Agriculture Organisation – United Nations agency, set

up in 1945, whose mandate is: ‘to raise levels of nutrition, improveagricultural productivity, better the lives of rural populations andcontribute to the growth of the world economy.’ Its annual $750million budget covers both ongoing programmes and emergencyrelief work

Farm-scale

evaluations

a $9 million research exercise in the UK to determine the on-farmeffects to fauna and flora of growing and managing herbicidetolerant crops compared to non-tolerant varieties of the samecrops

FMD foot and mouth disease – also known as hoof and mouth disease in

the USA, this virulent viral disease spread across the UK in 2001.Following scientific advice that has since been questioned, the UKgovernment implemented a drastic cull that resulted in theslaughter of 6 million animals, at an estimated cost to theeconomy of $15 billion

FSA Food Standards Agency (UK) – established in 2000 ‘to protect the

public’s health and consumer interests in relation to food.’Germplasm the genetic material, i.e the DNA, of an organism The term is

often used in connection with the collection or conservation ofseeds, cuttings, cell cultures, or other germplasm resources, inrepositories such as gene banks

GM genetically modified or genetically manipulated – a term normally

used to describe an organism into which DNA, containing one ormore genes, has been transferred from elsewhere The transferredDNA is never itself actually from another organism, but may be

an exogenous copy of DNA (i.e from a different species)

Alternatively the transferred DNA may be an extra copy of anendogenous gene (i.e from the same species) Finally, thetransferred DNA may be completely synthetic and hence of non-biological origin An organism containing any of these categories

of introduced gene is called transgenic

Heterosis also called hybrid vigour, the phenomenon whereby a hybrid of

genetically distinct (but often inbred) parents is sometimes muchmore vigorous than either parent In crop terms, hybridsexhibiting heterosis can out-yield their parents by as much as30–40%

HRI Horticulture Research Institute (UK) – also called Horticulture

Research International, HRI is a former public sector plantscience research centre that was transferred to ownership of theUniversity of Warwick in 2005

Hybrid an organism resulting from a cross between parents of differing

genotypes Hybrids may be fertile or sterile, depending onqualitative and/or quantitative differences in the genomes of thetwo parents Hybrids are most commonly formed by sexual cross-fertilisation between compatible organisms, but cell fusion andtissue culture techniques now allow their production from lessrelated organisms

ICARDA International Center for Agricultural Research in the Dry Areas –

this CGIAR-affiliated centre, established in 1977 with its quarters in Aleppo, Syria, has a mission ‘to improve the welfare ofpoor people and alleviate poverty through research and training indry areas of the developing world, by increasing the production,productivity and nutritional quality of food, while preserving andenhancing the natural resource base.’

improvement centres

ICRISAT International Crops Research Institute for the Semi-Arid Tropics

(India) – one of the CGIAR crop improvement centres

IFPRI International Food Policy Research Institute (USA) – one of the

CGIARagricultural improvement centres

IGER Institute for Grassland and Environmental Research (UK –

formed by a merger between the Welsh Plant Breeding Stationand the Grassland Research Institute at Hurley

IITA International Institute of Tropical Agriculture (Nigeria) – one of

the CGIAR crop improvement centres

ILRI International Livestock Research Institute (Kenya) – one of the

CGIARcrop improvement centres

Input trait a genetic character that affects how the crop is grown without

changing the nature of the harvested product For example,herbicide tolerance and insect resistance are agronomically usefulinput traits in the context of crop management, but they do notnormally alter seed quality or other so-called output traits that arerelated to the useful product of the crop

In situ

conservation

the maintenance of a species or population in its normal biologicalhabitat In the case of plants, this applies particularly to naturalpopulations of crop species and/or their wild relatives that may befuture sources of genetic variation, as well as to endangered species

in general In situ conservation is especially useful in thepreservation of traditional crop landraces, many of which areunder threat from the increasing use of higher yielding but moregenetically uniform modern varieties in agriculture

IPGRI International Plant Genetic Resources Institute (Italy) – an

international CGIAR-affiliated research institute with a mandate

to advance the conservation and use of genetic diversity for thewell being of present and future generations

Organization: ‘Intellectual property rights are the rights given topersons over the creations of their minds They usually give thecreator an exclusive right over the use of his/her creation for acertain period of time.’ IPR covers literary and artistic works (viacopyright) in addition to industrial inventions (via patents andtrademarks) and typically lasts for about 20 years IPR protection

of living organisms, such as plant varieties, is a more recent andcontroversial development

IRRI International Rice Research Institute (Philippines) – an

indepen-dent, non-profit agricultural research and training organisationand CGIAR centre that is focused on rice improvement IRRI wasestablished in 1960 by the Ford and Rockefeller foundations incooperation with the Philippines government with its main site atLos Ban˜os, near Manila

ISNAR International Service for National Agricultural Research (USA) –

assists developing countries in improving the performance of theirnational agricultural research systems and organisations bypromoting appropriate agricultural research policies, sustainableresearch institutions, and improved research management.IWMU International Water Management Institute (Sri Lanka) – one of

the CGIAR crop improvement centres

science research centre near Norwich

Land Grant

Universities

US network of agriculturally focused universities established bythe Morill Act in 1862

Landrace a genetically diverse and dynamic population of a given crop

produced by traditional breeding Landraces largely fell out offavour in commercial farming during the twentieth century andmany have died out Landraces are often seen as potentially usefulsources of novel genetic variation and efforts are under way toconserve the survivors

government department responsible for oversight of UK ture, including the commissioning of some research areas MAFFwas reorganised as DEFRA in 2001

PBIin the UK

Nutraceutical a neologism combining nutritional with pharmaceutical and

mean-ing a food product that has been determined to have a specificphysiological benefit for human health The term has no regulatorydefinition and is primarily used in promotion and marketing

of a consortium who agree to use the technologies in a particularmanner, e.g solely as non-profit public goods The best knownOAT is the Linux computer operating system, but analogousOATs have recently been developed in the field of agbiotech, mostnotably by CAMBIA.

Output trait a genetic character that alters the quality of the crop product itself,

e.g by altering its starch, protein, vitamin or oil composition.PBI Plant Breeding Institute – widely regarded as the premier centre of

plant breeding research in the UK, based in Cambridge, PBI wasprivatised in 1989 and subsequently sold on to a series ofmultinational companies

PBR Plant breeders’ rights – a form of intellectual property protection

in the European Union (via UPOV) designed specifically for newvarieties of plants

PCR Polymerase chain reaction – a technique for rapidly copying a

particular piece of DNA in the test tube (rather than in livingcells) PCR has made possible the detection of tiny amounts ofspecific DNA sequences in complex mixtures It is now used forDNA fingerprinting in police work, in genetic testing and in plantand animal breeding

PSIPRA Public Sector Intellectual Property Resource for Agriculture

(USA) – initiative of the Rockefeller and McKnight tions, in collaboration with ten of the major US LandGrant Universities As with CAMBIA, this US initiative isdesigned to support plant biotechnology research in developingcountries

Founda-PVPA Plant Variety Protection Act – legislation enacted in 1970 by the

US Congress that extended UPOV-like legal protection to plantgermplasm

Quantitative

genetics

the study of continuous traits (such as height or weight) and itsunderlying mechanisms

university research on the basis of the perceived quality of a ‘unit

of assessment’ that normally corresponds to a department Thisranking is then used to apportion funding selectively in favour ofhigher ranked departments

SAES State Agricultural Experiment Station (USA) – established by the

Hatch Act of 1887, the nationwide network of SAESs works withLand Grant Universities to carry out a joint research/teaching/extension mission

Species a group of organisms capable of interbreeding freely with each

other but not with members of other species (this is a muchsimplified definition, the species concept is much more complex) Aspecies can also be defined as a taxonomic rank below a genus,consisting of similar individuals capable of exchanging genes orinterbreeding

Teosinte the original wild grass, native to Mexico, from which cultivated

maize is derived; it is now classified as part of the same species asmaize, Zea mays

TILLING Targeting Induced Local Lesions IN Genomes – the directed

identification of random mutations controlling a wide range ofplant characters A more sophisticated DNA-based version ofmutagenesis breeding, TILLING does not involve transgenesis.Transgenesis the process of creating a transgenic organism

Transgenic an organism into which DNA, normally containing one or more

genes, has been transferred from elsewhere (see GM)

UPOV Union for the Protection of New Varieties – established in 1960 by

six European nations to extend legal ownership rights to plantgermplasm

USDA United States Department of Agriculture (USA) – established by

President Lincoln in 1862, USDA is the government departmentresponsible for all matters pertaining to agriculture, includingaspects of trade policy, food safety and the environment

WARDA Africa Rice Center (formerly called West Africa Rice

Develop-ment Association) – one of the CGIAR crop improveDevelop-ment centres

improvement centres

Wide crossing in plant breeding this refers to a genetic cross where one parent is

from outside the immediate gene pool of the other, e.g a wildrelative from one species crossed with a modern crop cultivar ofanother species

Wild relative plant or animal species that is taxonomically related to crop or

livestock species and serves as a potential source of genes forbreeding new crops or livestock varieties

The purpose of this book is to examine the wider scientific and social contexts

of modern plant breeding and agriculture We will begin by examining the historicaldevelopment of plant breeding over the past two centuries, before focusing on thedramatic changes of the last two decades Perhaps the best-known recent develop-ment in plant breeding is the emergence of genetic engineering, with its attendantsocial and scientific controversies But, as we shall see, GM crops and ‘agbiotech’(agricultural biotechnology) are just one manifestation of a more extensive series ofseismic changes that have profoundly altered the course of plant breeding since the1980s Today, in the middle of the first decade of the twenty-first century, plantbreeding and crop improvement are at an historic crossroads On one hand, are thetried and tested breeding methods that underpinned the Green Revolution andenabled us to feed the expanding world populations in the twentieth century Morerecently, however, governments across the world have largely dismantled theirapplied research infrastructures and have greatly reduced the capacity for public-good applications of newly emerging breeding technologies, including transgenesis.Much of this institutional restructuring occurred as part of the ideologically drivenprivatisation of public assets in the 1980s and 1990s The resulting depletion ofpublic sector breeding has left a void that was filled by a few private sector com-panies who applied a new paradigm of crop improvement based on transgenesis –and from this, the agbiotech revolution was born

As we confront the challenges of increasing populations, economic growth, risingaffluence, the spread of environmental degradation, and the depletion of non-renewable resources, twenty-first century agriculture will need all the tools andscientific expertise that plant breeders can muster Not to mention the appropriatecrop management strategies, market freedoms, and social stability that will benecessary to translate the promise of the breeder into the reality of productive andprofitable crops for the wellbeing of the farmer We will see how research into plantscience is becoming increasingly remote from its application for breeding For

1

a variety of different but linked reasons, public sector scientists are largely failing

to provide the requisite leadership in the development of practical public-goodtechnologies for crop improvement, especially in developing countries where theneed is greatest One of the main take-home messages of this book is that we must re-engage plant and agricultural science with the rest of society at a whole series oflevels These include better links between basic science and applied technologies,between scientific breeders and their farmer-customers, between the public sectorand the private sector, between industrialised countries and the developing world,between inexpensive conventional breeding and the costliest high-tech methods, andbetween agronomists and managers, and the economists and politicians working inagriculturally related areas of their respective professions

The book is divided into six parts that first introduce us to the science of plantbreeding before describing its changing social organisation and evolution as a mixedpublic/private venture over the last two centuries Part I includes a brief account ofthe origins of breeding and its transition from a farm-based empirical activity to thehighly sophisticated scientific programmes of today We will follow the increasinglysuccessful efforts of plant scientists of the eighteenth and nineteenth centuries toharness their growing knowledge of plant reproduction and development for prac-tical and profitable commercial application We will see how agricultural innovatorsbecame ever more skilled in manipulating those twin pillars of breeding, namelygenetic variation and selection The rediscovery of the principles of Mendelianinheritance and their application to simple and complex genetic traits was the keyscientific foundation of twentieth century crop breeding

The practical application of genetic knowledge to crop improvement in the field wasmade feasible by the theoretical and statistical tools provided by quantitative geneticsafter 1918 In the 1920s, chemical and X-ray mutagenesis were first used to create newcrop varieties, while the 1930s saw the beginnings of increasingly successful applica-tions of tissue culture in breeding programmes Soon, scientific breeders could createartificial hybrid combinations from different species, and even different genera And itwas not long before the first manmade crop species, a plant called triticale, wasproduced By the 1950s, the technique of wide crossing, coupled with chemicallyinduced chromosome manipulations, had enabled breeders to transfer chromosomes,

or parts thereof, from plants that were normally much too distantly related to breed More effective types of radiation mutagenesis, using nuclear sources such ascobalt-60 or caesium-137, were effectively used after World War II to create morethan 3000 new crop varieties

inter-In Part II, we will switch to consider the societal contexts of these scientificdevelopments that led us from the farmer-breeder of the nineteenth century

to today’s multinational, high-tech agribusiness During the nineteenth century,

it was realised that the most effective method for applying scientific principles tocrop improvement was to establish a professional body of trained plant breeders andresearchers In many of the newly industrialising countries, this was achieved bydirect government action Without a doubt, the most comprehensive, effective, andenduring crop improvement network is that of the USA, as originally established bythe Morrill Act in 1862, during the depths of the Civil War The British establishment,

in contrast, took a distinctly more laissez-faire route to agricultural betterment.Here, there was a gradual evolution of a disparate group of mostly privately fundedresearch centres during the late nineteenth and early twentieth centuries It was insome of these British research centres that the application of the newly rediscoveredprinciples of Mendelian genetics first propelled crop science into a new era In theUSA, the huge potential of hybrid crops, in terms of both yield and profitability,began to be realised during the 1920s with the introduction of the high-yieldingmaize varieties that eventually spread across the continent and beyond For most ofthe twentieth century, plant breeding and crop science research were very muchconcentrated in the public sector, with major contributions from universities andspecialised crop-focused research centres

The success of this public sector based paradigm became ever more apparent asincreasingly sophisticated breeding technologies were developed These technologies,developed by public sector plant researchers as free public goods, were called upon

to resolve the worsening food crisis as populations in developing countries expandedrapidly during the 1960s The Green Revolution of the 1960s and 1970s was largelythe result of the focused application of such public-good plant breeding, assisted bysome US-based philanthropic foundations Thanks to the work of a few groups ofdedicated plant breeders, new high-yielding varieties of wheat and rice were devel-oped, just in time to head off the spectre of mass hunger that haunted the Indiansubcontinent and much of Eastern Asia The spectacular success of the GreenRevolution in much (but not all) of the developing world led to the establishment of

an international network of plant research and breeding centres, including such vitalresources as seed and germplasm banks

In Part III, we move on to consider the turbulent events of the late twentiethcentury and the surprisingly rapid unravelling of the hitherto successful public/private paradigm of plant breeding research and development The 1970s and early1980s marked the apogee of public sector and public-good international plantbreeding Within a few years, governments around the world began to dismantletheir public sector plant science infrastructures, in line with the new privatisationagenda that emanated largely from the UK Meanwhile, the private sector emergedfrom the shadows as an increasingly dominant force in the enterprise of cropmodification and improvement Two additional factors facilitated the growth of the

private sector: the shift to a more benign regulatory environment for the legalprotection of new plant varieties; and the invention of a new set of plant manip-ulation technologies that would allow the patenting of transgenic (GM) crop vari-eties We will go on to follow the fate of some of the rationalised, reduced, orterminated public breeding programmes across the world and the resulting retreat ofthe vast majority of public sector researchers into more academic studies.

The topic of Part IV is agbiotech The decade from 1985–1995 witnessed a damental shift in the world of plant breeding, as the private sector became the moredominant partner and transgenic technologies were increasingly promoted as theway forward for crop improvement in general We will analyse the consequences ofthese important developments for the future of agriculture I will present the casethat it was not so much genetic engineering (transgenesis) itself that has been the rootcause of the many public controversies about agricultural biotechnology (agbiotech).Rather, it is the context in which the technology was created, promoted, and thenapplied to crop manipulation, which was radically different to previous forms ofhigh-tech scientific crop improvement After World War II, highly intrusive and

fun-‘artificial’ methods of crop genetic modification had already been developed in thepublic domain with little or no fanfare or public controversy These technologieswere used freely to create new crop varieties around the world and were especiallywidely applied in developing countries

In contrast, transgenic technologies were largely developed and patented by theprivate sector Some companies then used the new technologies for the manipulation

of a few simple input traits in a few profitable commercial cash crops In themeantime, however, these technologies had already been widely hailed, by publicsector scientists and companies alike, as a radical and revolutionary breakthrough inplant breeding of almost unlimited potential for the future of agriculture Subse-quently, the fact that, notwithstanding the optimistic rhetoric, nothing of anymatching public value has so far emerged from transgenesis, has engendered

a mixture of public scepticism and distrust about the entire agbiotech enterprise Wewill also see how the actions of a few agbiotech companies are currently in danger ofsabotaging some rather promising future developments in transgene technology toproduce cheap medicines via biopharming

In Part V, we will discuss alternative methods of enhancing crop production,especially amongst the rapidly increasing populations of the developing world I willshow that there need not be any looming crisis in feeding the world population overthe next fifty years We already have the crops, the breeding expertise, and theorganisational skills to achieve this task – providing it is managed properly I willpresent the case for a judicious expansion of our use of arable land, especially inparts of South America, where a large amount of non-forested land is available for

sustainable crop cultivation Combined with re-use of fallow, abandoned, andset-aside land, these measures could significantly increase global food productionover the next few decades Other productivity enhancing measures include better on-farm management, improvement of physical and regulatory infrastructure (ports,roads, credit facilities, tax regimes etc.), and the ending of discriminatory tariffs andsubsidies Implementation of these exceedingly practical but relatively unglamorousmeasures, along with the prospect of continuing yield gains via plant breeding,should ensure that we will be able to ‘feed the world’ over the next fifty years,without recourse to more nebulous and uncertain ‘magic bullet’ solutions.

In Part VI, we will look forward to the future of plant breeding in the twenty-firstcentury, whether in the public/private sectors, or in industrial/developing countries

We will discuss the uncertain situation of international organisations like CGIAR(Consultative Group on International Agricultural Research), our endangered glo-bal seed banks, and the often heroic, and largely unseen, efforts of breeders incountries from Iraq to Coˆte d’Ivoire in trying to maintain these precious resourcesagainst the depredations of warfare and civil strife and the more benign neglect ofincreasingly jaded funding bodies We will then look forward to consider some newoptions that could allow a reinvigorated public sector to resume its place as a majorpartner in the global enterprise of crop improvement The long-term success ofinternational agriculture is dependent on a diverse, mixed ecology of public andprivate agents and agencies We need strong, well-resourced public-good ventures,which in turn are balanced and complemented by appropriately regulated, for-profit,private sector ventures that are both innovative and truly competitive

The current problems of plant breeding have not been helped by the fact thatmany public sector scientists have largely withdrawn from practical breeding andpublic debate, to the more secluded and serene realms of basic research The latterare not so much ivory towers as ivory cloisters of an almost adamantine unworld-liness This withdrawal has left the public arena bereft of many of the voices thatcould bring some balance into the sterile and polarised discourse on transgenic cropsthat has plagued the debates of the past decade It is only by regaining a sense ofbalance in each of these aspects of crop improvement that we can recapture publicconfidence, and move forward with a renewed sense of optimism to confront andresolve the many challenges of agriculture in the twenty-first century

Part I The science of plant breeding

Here Ceres’ gifts in waving prospect stand,And nodding tempt the joyful reaper’s hand

Alexander Pope (1688–1744)Windsor Forest(l 39)

1 Origins of plant breeding

For out of olde feldes, as men seithCometh al this newe corn fro yeer to yere;

And out of olde bokes, in good feith,Cometh all this newe science that men lere

Geoffrey Chaucer (c 1382)The Parlement of FoulesIntroduction – the development of agricultureFor most of our history, we humans have been omnivores who enjoyed a variedplant and animal based diet that was derived from a hunter-gathering lifestyle Thisspecial relationship has bound people and plants in mutual dependence for well overone hundred millennia During this period, our Palaeolithic and Neolithic ancestorsexperimented with many different strategies of plant exploitation, especially duringthe last hundred millennia when climatic conditions changed repeatedly and otherresources such as large animals often became progressively more difficult to obtain.1For tens of millennia before the start of formal agriculture, societies throughout theworld were engaged in many types of relatively sophisticated management of theirfavoured food plants For example, 23 000 years ago, people in the Jordan Valleywere already harvesting and grinding wild cereal grains, and baking the flour intobread and cakes.2Discoveries of similar grinding implements dating back as far as

48 000 years ago might mean that the management and processing of cereals went onfor well over 30 000 years before these plants were ever cultivated as crops

During this period of informal plant management, our ancestors unwittinglybegan a process of plant selection that would lead to the domestication of a fewgenetically amenable species These plants became the first successful crops, and theirformal cultivation was already well under way in several regions of the world by

12 000 years ago, and possibly earlier Following the development and spread of

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agriculture, most of our ancestors came to rely increasingly on a much morerestricted repertoire of domesticated plants for the majority of their food needs Eventoday, most of the world depends on a small, carefully selected group of edibleplants We also use plants for a host of other purposes, such as clothing, shelter,medicines and tools Although we now believe that the beginnings of the domes-tication process were probably non-intentional and unforeseen by Palaeolithic andNeolithic proto-farmers, these people soon learned how to improve their new crops

by conscious forms of selection and breeding.3

Non-intentional selectionSelection can be said to be the backbone of crop breeding There is little point inassembling or creating a group of genetic variants unless one has an effectivemechanism to recognise and select the best adapted or most useful of these variants forfurther propagation Such selection could have been either unintentional or deliberate

on the part of the early farmers or would-be farmers To a great extent, all livingorganisms act in concert with the abiotic (non-living) part of the environment asunintentional agents of selection This sort of selection is normally negative, i.e less fitindividuals tend to be eliminated from the population For example, simply by huntingfor prey, a carnivore will tend to select those individuals that are easier to capturebecause they are less well protected, slower, more easily detected etc As a result, theprey population is selected in favour of fitter individuals who, for example, may haveadopted herding behaviour, are more fleet of foot, have acquired camouflage etc Ourancestors started out as non-intentional agents of selection during the early stages ofcrop domestication They then progressed to conscious selection of the relatively smallnumber of favourable traits that were readily recognisable in a crop plant, e.g seedsize, vigour, or yield In contrast, the scientific breeders of today have access to abattery of screening and selection strategies, many of them automated, that enablethem to manipulate hundreds of often invisible traits in our major crop plants.Most plants in non-agricultural ecosystems have been selected for traits such asindeterminate flowering and easy seed shedding from the parent plant This mini-mises the chance of seed loss to herbivores Several early human societies usedtechniques of plant management that had the by-product of selecting for a differentset of traits From recent genetic evidence, we know that these new traits enabled afew plant species to develop in the direction of domestication The concept of non-intentional, or unconscious, selection by humans was first expounded by Darwin,4although he had no idea of the mechanism by which the favoured variants couldtransmit their variations to subsequent generations.5More recently, this mechanism

of crop selection has been described in detail by Zohary.6Non-intentional selection