Cereal biotechnology Edited by Peter C Morris and James H Bryce Published byWoodhead Publishing Limited Abington Hall, Abington Cambridge CB1 6AH England Published in Northand SouthAmerica byCRCPressLLC 2000Corporate Blvd, NW Boca Raton FL33431 USA First published 2000,Woodhead Publishing Limited and CRCPressLLC (cid:1) 2000, Woodhead Publishing Limited The authors have asserted their moral rights. Conditions of sale This bookcontains information obtained from authentic andhighly regarded sources. Reprinted material isquoted with permission, and sourcesare indicated. Reasonable efforts have been made to publish reliable data and information, but the authors and the publisherscannot assume responsibility forthe validity ofall materials. Neither the authors nor the publishers, noranyone else associated withthis publication, shall be liable for anyloss,damage orliability directly or indirectly caused or alleged to be caused bythisbook. 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Woodhead Publishing Limited ISBN1 855734982 CRCPressISBN0849308992 CRCPressorder number: WP0899 Cover design byThe ColourStudio Project managed byMacfarlane Production Services, Markyate, Hertfordshire Typeset byMHLTypesetting Limited, Coventry Printed byT JInternational, Padstow,Cornwall, England Related titles from Woodhead’s food science, technology and nutrition list: Kent’s technology of cereals (ISBN:1 855733617) NL Kentand ADEvers This well established textbook provides an authoritative andcomprehensive study of cereal technology. Food machinery (ISBN:1 855732696) L MCheng This bookprovides ageneral technical and mechanical background forthe basic processing machinery nowusedfor making snacks,baked goodsandconfectionery. It covers the basic principles, machine design, function, operation and output. Wheat – Chemistryandutilization (ISBN:1 566763487) HJCornell and AW Hoveling This bookprovides the reader with extensive newinformation onwheat components that will beuseful in improving utilization of wheat and the formulation of newand up-graded wheat-based foodproducts. Details ofthese booksand acomplete list of Woodhead’s food science, technology and nutrition titles can beobtained by: • visiting our website at www.woodhead-publishing.com • contacting Customer Services (e-mail: [email protected]; fax: +44 (0)1223 893694;tel: +44(0)1223 891358ext. 30; address:Woodhead Publishing Ltd, Abington Hall, Abington, Cambridge CB1 6AH,England) If youwouldlike to receive information onforthcoming titles in this area, please send your addressdetails to: Francis Dodds(address,tel. and fax as above; e-mail: [email protected]).Please confirm which subject areas youare interested in. Contents 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 vi 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 Contents vii 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 viii Contents 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 Contributors Chapters 1 and 11 Chapter 3 Dr Peter Morris and Dr James Bryce Dr M R Davey Department of Biological Sciences Faculty of Science Heriot-Watt University University Park Riccarton Nottingham NG7 2RD Edinburgh EH14 4AS Tel: +44 (0) 115 951 3057 Scotland Fax: +44 (0) 115 951 3298 Tel: +44 (0) 131 451 3181 E-mail: [email protected] Fax: +44 (0) 131 451 3009 E-mail: [email protected] Chapter 4 Dr David McElroy Director of Agricultural Business Chapter 2 Operations Dr R Schuurink and Ms J Louwerse Maxygen Inc Department of Biological Sciences 515 Galveston Drive Heriot-Watt University Redwood City Riccarton CA 94063 Edinburgh EH14 4AS USA Scotland Tel: +1 650 298 5454 Tel: +44 (0) 131 449 5111 Fax: +1 650 364 2715 Fax: +44 (0) 131 451 3009 E-mail: [email protected] x Contributors Chapter 5 Chapter 8 Professor Robert Henry Dr Andrew Lynn Centre forPlantConservationGenetics Food Standards & Product Southern Cross University Technology Department PO Box 157 SAC Lismore Auchincruive NSW Ayr KA6 5HW AUSTRALIA Scotland Tel: (02) 6620 3010 Tel: +44 (0) 1292 525087 Fax: (02) 6622 2080 Fax: +44 (0) 1292 525071 E-mail: [email protected] E-mail: [email protected] Chapter 6 Chapter 9 Dr Bill Thomas Dr Ray Anderson Scottish Crop Research Institute High Ridge Invergowrie Marchington Dundee DD2 5DA Uttoxeter Scotland Staffs ST14 8LH Tel: +44 (0) 1382 562731 Tel: +44 (0) 1283 820333 Fax: +44 (0) 1382 562426 E-mail: E-mail: [email protected] [email protected] Chapter 7 Chapter 10 Dr Wendy Cooper Dr Eric Evans PO Box 686 Department of Agriculture Norwich NR5 0PZ University of Newcastle Newcastle upon Tyne NE1 7RU Tel: +44 (0) 1603 741293 E-mail: [email protected] Tel: +44 (0) 191 222 6925 Fax: +44 (0) 191 222 7811 Dr Jermy Sweet E-mail: [email protected] National Institute of Agricultural Botany Huntingdon Road Cambridge CB3 0LE Tel: +44 (0) 1223 276381 Fax: +44 (0) 1223 277602 1 Introduction P. C. Morris and J. H. Bryce, Heriot-Watt University, Edinburgh 1.1 Cereals: an introduction CerealsowetheirEnglishnametotheRomangoddessCeres,thegiverofgrain, 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 indirectlyasanimalfeed.Itisprimarilythegrainsofcerealsthatareusefultous, 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.Thefloweringorgansarecarriedonastemcalledtherachis,which may be branched, and in turn bears spikelets which may carry more than one flowerateachnodeoftherachilla(Fig.1.1).Thespikeletsmaybeorganisedina 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,andeach flower inthe spikelet isenclosedintwobract-like organscalledthelemmaandpalea.Thelemmamaybeextendedtoformalong awn. In some cereals or cereal varieties the lemma and palea may remain attachedtothegrain;thesearetermedhulledorhuskedgrains,suchasoatsand 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