cereal biotechnology

Cereal biotechnology Edited by Peter C Morris and James H Bryce Published by Woodhead Publishing Limited Abington Hall, Abington Cambridge CB1 6AH England Published in North and South America by CRC Press LLC 2000 Corporate Blvd, NW Boca Raton FL 33431 USA First published 2000, Woodhead Publishing Limited and CRC Press LLC ß 2000, Woodhead Publishing Limited The authors have asserted their moral rights. Conditions of sale This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. Reasonable efforts have been made to publish reliable data and information, but the authors and the publishers cannot assume responsibility for the validity of all materials. Neither the authors nor the publishers, nor anyone else associated with this publication, shall be liable for any loss, damage or liability directly or indirectly caused or alleged to be caused by this book. 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Woodhead Publishing Limited ISBN 1 85573 498 2 CRC Press ISBN 0 8493 0899 2 CRC Press order number: WP0899 Cover design by The ColourStudio Project managed by Macfarlane Production Services, Markyate, Hertfordshire Typeset by MHL Typesetting Limited, Coventry Printed by T J International, Padstow, Cornwall, England Related titles from Woodhead’s food science, technology and nutrition list: Kent’s technology of cereals (ISBN: 1 85573 361 7) N L Kent and A D Evers This well established textbook provides an authoritative and comprehensive study of cereal technology. Food machinery (ISBN: 1 85573 269 6) L M Cheng This book provides a general technical and mechanical background for the basic processing machinery now used for making snacks, baked goods and confectionery. It covers the basic principles, machine design, function, operation and output. Wheat – Chemistry and utilization (ISBN: 1 56676 348 7) H J Cornell and A W Hoveling This book provides the reader with extensive new information on wheat components that will be useful in improving utilization of wheat and the formulation of new and up-graded wheat-based food products. Details of these books and a complete list of Woodhead’s food science, technology and nutrition titles can be obtained by: • visiting our web site at www.woodhead-publishing.com • contacting Customer Services (e-mail: sales@woodhead-publishing.com; fax: +44 (0)1223 893694; tel: +44 (0)1223 891358 ext. 30; address: Woodhead Publishing Ltd, Abington Hall, Abington, Cambridge CB1 6AH, England) If you would like to receive information on forthcoming titles in this area, please send your address details to: Francis Dodds (address, tel. and fax as above; e-mail: francisd@woodhead-publishing.com). Please confirm which subject areas you are interested in. List of contributors ix 1 Introduction 1 P. C. Morris and J. H. Bryce, Heriot-Watt University, Edinburgh 1.1 Cereals: an introduction 1 1.2 Plant breeding 8 1.3 Biotechnology: an introduction 13 1.4 The structure of this book . . . . 14 1.5 Sources of further information and advice 14 1.6 References 15 2 The genetic transformation of wheat and barley 17 R. C. Schuurink and J. D. Louwerse, Heriot-Watt University, Edinburgh 2.1 Introduction 17 2.2 Issues in successful transformation 19 2.3 Target tissues for transformation . . . 20 2.4 Delivery of DNA 25 2.5 Selection and regeneration 28 2.6 Promoters 31 2.7 Examples of transformed wheat and barley . 33 2.8 Summary: problems and future trends 35 2.9 Sources of further information and advice 36 2.10 References 37 Contents 3 The genetic transformation of rice and maize 43 M. R. Davey, H. Ingram, K. Azhakanandam and J. B. Power, University of Nottingham 3.1 Introduction 43 3.2 Approaches to the transformation of maize and rice 44 3.3 Target tissues for rice and maize transformation . . . . . . . 50 3.4 Vectors for rice and maize transformation 52 3.5 Examples of agronomically useful genes introduced into rice and maize 55 3.6 Summary: problems, limitations and future trends 57 3.7 Sources of further information and advice 60 3.8 Internet sites . . . 61 3.9 References 61 4 Product development in cereal biotechnology 71 D. McElroy, Maxygen Inc., Redwood City 4.1 Introduction 71 4.2 Commercial targets for cereal biotechnology . 72 4.3 Problems in cereal biotechnology . . . 74 4.4 Efficacy screening of commercial traits 75 4.5 Molecular breeding of transgenic plants 78 4.6 Molecular quality control for transgenic plants . . . . . . 79 4.7 Intellectual property and freedom to operate 81 4.8 Regulatory issues and risk assessment 82 4.9 Product release and marketing strategies 84 4.10 Product development: a practical example 84 4.11 Future trends 88 4.12 References 89 5 Using biotechnology to add value to cereals 91 R. J. Henry, Southern Cross University, Lismore 5.1 Introduction 91 5.2 Weed control (productivity, quality, safety) 92 5.3 Disease resistance (productivity, quality, safety) 93 5.4 Improved nutritional properties (quality, safety) . . . . . . 95 5.5 Improved processing properties (productivity, quality, safety) 96 5.6 Improved cereal quality control (quality, safety) . . 103 5.7 Summary: future prospects and limitations 103 5.8 Sources of further information and advice 104 5.9 References 105 6 Molecular biological tools in cereal breeding 107 W. Thomas, Scottish Crop Research Institute, Dundee 6.1 Introduction 107 vi Contents 6.2 Markers 110 6.3 Characters . . . 116 6.4 Deployment of molecular markers 119 6.5 Future prospects . . . . . . . 124 6.6 Conclusions 126 6.7 Sources of further information and advice 126 6.8 References 127 7 Risk assessment and legislative issues 137 W. Cooper, formerly NIAB, Cambridge and J. B. Sweet, NIAB, Cambridge 7.1 Introduction 137 7.2 Risk assessment and avoidance: general principles 140 7.3 Assessing the impact of genetically modified crops 145 7.4 How is biotechnology regulated? 149 7.5 Public perceptions . . . 154 7.6 Future developments in the regulatory process . . . 157 7.7 References 158 8 Current practice in milling and baking 161 A. Lynn, Scottish Agricultural College, Auchincruive 8.1 Introduction 161 8.2 Composition of cereals . 162 8.3 Use of cereals in milling 166 8.4 Cereal requirements for milling 169 8.5 Use of cereals in baking 171 8.6 Bread baking . 172 8.7 Biscuit manufacture . . . . 175 8.8 Summary 175 8.9 Bibliography . 178 8.10 References 180 9 Current practice in malting, brewing and distilling 183 R. G. Anderson, Marchington Zymoscience, Uttoxeter 9.1 Introduction 183 9.2 Fundamentals of malting, brewing and distilling . . . 184 9.3 Malting industry: current practice 186 9.4 Brewing industry: current practice 194 9.5 Distilling industry: current practice 200 9.6 Summary: limitations in current practice and the role of biotechnology 207 9.7 Sources of further information and advice 209 9.8 References 211 Contents vii 10 Current practice in cereal production 217 E. J. Evans, University of Newcastle 10.1 Introduction 217 10.2 Varietal selection 227 10.3 Crop establishment . . . . . . 228 10.4 Crop nutrition 229 10.5 Weed control 232 10.6 Disease control 233 10.7 Pest control . . . . . . . 234 10.8 Harvesting and grain storage . . . 235 10.9 References and further reading 236 11 Summary and conclusions 237 P. C. Morris, G. H. Palmer and J. H. Bryce, Heriot-Watt University, Edinburgh 11.1 Improving cereal production and quality: a global challenge 237 11.2 Potential of cereal biotechnology 238 11.3 Biotechnology in commercial practice 239 11.4 Problems facing the cereal biotechnology industry . . . . . 240 11.5 The future 241 Index . . . . . . . . . . . . . 243 viii Contents Chapters 1 and 11 Dr Peter Morris and Dr James Bryce Department of Biological Sciences Heriot-Watt University Riccarton Edinburgh EH14 4AS Scotland Tel: +44 (0) 131 451 3181 Fax: +44 (0) 131 451 3009 E-mail: P.C.Morris@hw.ac.uk Chapter 2 Dr R Schuurink and Ms J Louwerse Department of Biological Sciences Heriot-Watt University Riccarton Edinburgh EH14 4AS Scotland Tel: +44 (0) 131 449 5111 Fax: +44 (0) 131 451 3009 Chapter 3 Dr M R Davey Faculty of Science University Park Nottingham NG7 2RD Tel: +44 (0) 115 951 3057 Fax: +44 (0) 115 951 3298 E-mail: mike.davey@nottingham.ac.uk Chapter 4 Dr David McElroy Director of Agricultural Business Operations Maxygen Inc 515 Galveston Drive Redwood City CA 94063 USA Tel: +1 650 298 5454 Fax: +1 650 364 2715 E-mail: david_mcelroy@maxygen.com Contributors Chapter 5 Professor Robert Henry Centre for Plant Conservation Genetics Southern Cross University PO Box 157 Lismore NSW AUSTRALIA Tel: (02) 6620 3010 Fax: (02) 6622 2080 E-mail: rhenry@scu.edu.au Chapter 6 Dr Bill Thomas Scottish Crop Research Institute Invergowrie Dundee DD2 5DA Scotland Tel: +44 (0) 1382 562731 Fax: +44 (0) 1382 562426 E-mail: wthoma@scri.sari.ac.uk Chapter 7 Dr Wendy Cooper PO Box 686 Norwich NR5 0PZ Tel: +44 (0) 1603 741293 E-mail: wendy.cooper@ukgateway.net Dr Jermy Sweet National Institute of Agricultural Botany Huntingdon Road Cambridge CB3 0LE Tel: +44 (0) 1223 276381 Fax: +44 (0) 1223 277602 Chapter 8 Dr Andrew Lynn Food Standards & Product Technology Department SAC Auchincruive Ayr KA6 5HW Scotland Tel: +44 (0) 1292 525087 Fax: +44 (0) 1292 525071 E-mail: A.Lynn@au.sac.ac.uk Chapter 9 Dr Ray Anderson High Ridge Marchington Uttoxeter Staffs ST14 8LH Tel: +44 (0) 1283 820333 E-mail: raymond.anderson@care4free.net Chapter 10 Dr Eric Evans Department of Agriculture University of Newcastle Newcastle upon Tyne NE1 7RU Tel: +44 (0) 191 222 6925 Fax: +44 (0) 191 222 7811 E-mail: E.J.Evans@ncl.ac.uk x Contributors 1.1 Cereals: an introduction Cereals owe their English name to the Roman goddess Ceres, the giver of grain, indicative of the antiquity and importance of cereals (Hill 1937). This importance is still very much the case today; cereals of one sort or another sustain the bulk of mankind’s basic nutritional needs, both directly and indirectly as animal feed. It is primarily the grains of cereals that are useful to us, although the vegetative parts of the plant may be used as fodder or for silage production, and straw is used for animal bedding. Cereals are members of the large monocotyledonous grass family, the Gramineae. The flowering organs are carried on a stem called the rachis, which may be branched, and in turn bears spikelets which may carry more than one flower at each node of the rachilla (Fig. 1.1). The spikelets may be organised in a loose panicle as in sorghum, oats and some millets, or in a tight spike, as in wheat. The length of the internodes of the rachis and of the rachilla, and the number of flowers at each node of the spikelet determine the overall architecture. Each spikelet is subtended by two bracts or leaf-like organs termed the glumes, and each flower in the spikelet is enclosed in two bract-like organs called the lemma and palea. The lemma may be extended to form a long awn. In some cereals or cereal varieties the lemma and palea may remain attached to the grain; these are termed hulled or husked grains, such as oats and most barleys, as opposed to naked grains such as most wheats and maize (Fig. 1.1). The cereals, with the exception of maize, are dioecious. Each flower bears both male organs; the three anthers (six in rice), and female organs; the ovary which carries two feathery stigmas. In maize, the male flowers are borne in 1 Introduction P. C. Morris and J. H. Bryce, Heriot-Watt University, Edinburgh [...]... order to spread their offspring far and wide In wild cereals, the spikes bearing the grains Introduction 9 Fig 1.4 Annual global cereal production in millions of tonnes and cereal yield in tonnes per hectare for the years 1961–98 (from FAO data) Fig 1.5 Annual global fertiliser use and pesticide imports over years 1961–97 (from FAO data) 10 Cereal biotechnology are borne on a brittle rachis (the main... baking, and in cereal production, three technology-intensive industries that work with cereals as their prime raw materials 1.5 Sources of further information and advice http://www.hgca.com/ The Home-Grown Cereals Authority exists to improve the production and marketing of UK cereals http://www.smallgrains.org/Index.htm A site focusing on the production and marketing of North American cereals Introduction... are troublesome weeds (for example, Avena fatua) Oats form a minor component of the world cereal crop at 1.5% of total global cereal production (Fig 1.2) and the biggest current producers are Russia, Canada and the USA (Fig 1.3) 1.1.6 Rye Rye (Secale cereale) appears to be more recently domesticated than other cereals, although it was known to the ancient Greek and Roman civilisations (DeCandolle 1886)...2 Cereal biotechnology Fig 1.1 Generalised structure of cereal flowering organs The length and branching pattern of the rachis and the rachilla, and the number of flowers per spikelet determine the overall appearance of the cereal spikes on a terminal panicle called a tassel, and the female flowers are in spikelets... yield than winter wheats (Peterson 1965, Pomeranz 1987) Wheat is one of the most widely grown cereals, accounting for over onequarter of the world’s global cereal production, and is primarily used for human consumption with some 15% being used for animal feed The largest global Fig 1.3 Continued 6 Cereal biotechnology producers of wheat are China, India and the USA (Fig 1.3) Wheats can be classified... stable integration and expression of genetic 18 Cereal biotechnology information which is introduced into wheat and barley by means other than breeding via crosses In other words, heterologous (derived from a different species) or modified homologous (derived from the same species) genes are introduced into the genetic blueprint (the genome) of the cereal The cereal will express this new genetic information... predominantly produced in central and eastern Europe Both rye and oats may have originated as weed 8 Cereal biotechnology species in wheat and barley crops Rye is a very winter-hardy crop that will grow on poor soils such as those of the north European plain Rye accounts for only around 1% of world total cereal production (Fig 1.2), is used for animal food and human consumption and the prime producers... still grown in some parts of Europe It is thought to be descended from the wild species, T dicoccoides, which is still found in the Fig 1.2 Annual global production of cereals in millions of tonnes (from FAO data for 1998) 4 Cereal biotechnology eastern Mediterranean region Triticum durum (macaroni wheat), in turn descended from emmer, is grown world wide and has excellent pasta-making qualities Triticum... throughout the world, as a source of genetic variation for the crops of tomorrow (Chrispeels and Sadava 1994) 1.3 Biotechnology: an introduction Biotechnology is a difficult term to define since the harnessing of any biological process to human aims and desires could justifiably be called biotechnology However, the revolution in our understanding of the molecular mechanisms underlying the processes of... DNA, the prime genetic material, has resulted in the ability to manipulate those mechanisms to our requirements This new-found knowledge and ability is loosely termed biotechnology There are two main applications of biotechnology to cereals The first is as an aid to conventional breeding programmes, as outlined above Physiological or morphological traits are governed by genes carried on chromosomes . Product development in cereal biotechnology 71 D. McElroy, Maxygen Inc., Redwood City 4.1 Introduction 71 4.2 Commercial targets for cereal biotechnology . 72 4.3 Problems in cereal biotechnology Improving cereal production and quality: a global challenge 237 11.2 Potential of cereal biotechnology 238 11.3 Biotechnology in commercial practice 239 11.4 Problems facing the cereal biotechnology. offspring far and wide. In wild cereals, the spikes bearing the grains 8 Cereal biotechnology Fig. 1.4 Annual global cereal production in millions of tonnes and cereal yield in tonnes per hectare