BIOLOGY OF BREEDING POULTRY Poultry Science Symposium Series Executive Editor (Volumes 1–18): B.M. Freeman 1 Physiology of the Domestic Fowl* 2 Protein Utilization by Poultry* 3 Environmental Control in Poultry Production* 4 Egg Quality – a Study of the Hen’s Egg* 5 The Fertility and Hatchability of the Hen’s Egg* 6 i. Factors Affecting Egg Grading* ii. Aspects of Poultry Behaviour* 7 Poultry Disease and World Economy 8 Egg Formation and Production 9 Energy Requirements of Poultry* 10 Economic Factors Affecting Egg Production* 11 Digestion in the Fowl* 12 Growth and Poultry Meat Production* 13 Avian Coccidiosis* 14 Food Intake Regulation in Poultry* 15 Meat Quality in Poultry and Game Birds 16 Avian Immunology 17 Reproductive Biology of Poultry 18 Poultry Genetics and Breeding 19 Nutrient Requirements of Poultry and Nutritional Research* 20 Egg Quality – Current Problems and Recent Advances* 21 Recent Advances in Turkey Science 22 Avian Incubation 23 Bone Biology and Skeletal Disorders 24 Poultry Immunology* 25 Poultry Meat Science 26 Poultry Feedstuffs 27 Welfare of the Laying Hen 28 Avian Gut Function in Health and Disease 29 Biology of Breeding Poultry *Out of print Volumes 1–24 were not published by CAB International. Those still in print may be ordered from: Carfax Publishing Company PO Box 25, Abingdon, Oxfordshire OX14 3UE, UK Biology of Breeding Poultry Poultry Science Symposium Series Volume Twenty-nine Edited by P.M. Hocking Division of Genetics and Genomics, The Roslin Institute and Royal (Dick) School of Veterinary Studies, University of Edinburgh, U K CABI is a trading name of CAB International CABI Head Office Nosworthy Way Wallingford Oxfordshire OX10 8DE UK Tel: +44 (0)1491 832111 Fax: +44 (0)1491 833508 E-mail: cabi@cabi.org Website: www.cabi.org CABI North American Office 875 Massachusetts Avenue 7th Floor Cambridge, MA 02139 USA Tel: +1 617 395 4056 Fax: +1 617 354 6875 E-mail: cabi-nao@cabi.org © CAB International 2009. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Biology of breeding poultry / edited by P.M. Hocking. p. cm. (Poultry science symposium series ; v. 29) Includes bibliographical references and index. ISBN 978-1-84593-375-3 (alk. paper) 1. Poultry Breeding Congresses. 2. Poultry Genetics Congresses. I. Hocking, P. M. (Paul M.) II. Title. III. Series: Poultry science symposium ; no. 29. SF492.B56 2009 636.5'082 dc22 2008033138 ISBN-13: 978 1 84593 375 3 Typeset by Columns Design Ltd, Reading. Printed and bound in the UK by the MPG Books Group, Bodmin. The paper used for the text pages in this book is FSC certified. The FSC (Forest Stewardship Council) is an international network to promote responsible management of the world's forests. v CO N T E N T S CO N T R I B U T O R S viii PR E F A C E xi AC K N O W L E D G E M E N T S xiii PA R T I Introduction 1 CH A P T E R 1 The Genetics of Modern Commercial Poultry 3 J.C. McKay CH A P T E R 2 Breeder Management: How Did We Get Here? 10 K.F. Laughlin PA R T II Genetic Improvement 27 CH A P T E R 3 Developments in Quantitative Genetics and Genomics Relevant for Poultry Breeding 29 P. Bijma and H. Bovenhuis CH A P T E R 4 Applications in Poultry Production 45 H.M. Sang CH A P T E R 5 Prospects for Sex Determination in Poultry 54 S. Nandi and M. Clinton vi Contents PA R T III Physiology of Reproduction 59 CH A P T E R 6 Endocrinology and Genetics of the Hypothalamic–Pituitary– Gonadal Axis 61 I.C. Dunn, N.A. Ciccone and N.T. Joseph CH A P T E R 7 Control of Follicular Development: Intra-ovarian Actions of Transforming Growth Factor-β (TGF-β) Superfamily Members 89 P.G. Knight, S.L. Al-Musawi, T.M. Lovell and R.T. Gladwell PART IV Mating Behaviour and Fertility 109 CH A P T E R 8 Mating Behaviour and Fertility 111 I.J.H. Duncan CH A P T E R 9 Sperm Competition and Fertilization Success 133 T.R. Birkhead and T. Pizzari CH A P T E R 10 Semen Quality and Semen Storage 151 G.J. Wishart PA R T V Incubation and Hatching 179 CH A P T E R 11 Broodiness and Broody Control 181 P.J. Sharp CH A P T E R 12 Incubation and Hatching 206 N.A. French CH A P T E R 13 V. Bruggeman, K. Tona, O. Onagbesan and E. Decuypere PA R T VI Managing the Environment 241 CH A P T E R 14 Photoperiod and Control of Breeding 243 P.D. Lewis Contents vii CH A P T E R 15 Behaviour and Environmental Enrichment in Broiler Breeders 261 I. Estevez CH A P T E R 16 Ratites, Game Birds and Minor Poultry Species 284 D.C. Deeming PA R T VII Nutrition of Breeding Poultry 305 CH A P T E R 17 Feed Restriction 307 P.M. Hocking CH A P T E R 18 Protein and Amino Acid Responses 331 C. Fisher and R.M. Gous CH A P T E R 19 Vitamins, Minerals and Micronutrients 361 M.T. Kidd PA R T VIII Health and Welfare 375 CH A P T E R 20 Vaccination: Theory and Practice 377 T. Cserep CH A P T E R 21 Immune Protection of the Hatchling 391 C. Butter and H.J. Walter CH A P T E R 22 Managing Current Disease Challenges in Breeders 414 S.R. Collett PA R T IX Abstracts 435 CH A P T E R 23 Poster Abstracts 437 Index 455 viii CO N T R I B U T O R S S.L. Al-Musawi, Department of Veterinary Basic Sciences, The Royal Veterinary College, Camden, London NW1 0TU; email: salmusawi@rvc. ac.uk P. Bijma, Animal Breeding and Genomics Centre, Wageningen University, Marijkeweg 40, 6709PG Wageningen, The Netherlands; email: piter. bijma@wur.nl T.R. Birkhead, Department of Animal and Plant Sciences, University of H. Bovenhuis, Animal Breeding and Genomics Centre, Wageningen University, Marijkeweg 40, 6709PG Wageningen, The Netherlands; email: henk.bovenhuis@wur.nl V. Bruggeman, K.U.Leuven, Faculty of Bioscience Engineering, Depart- ment of Biosystems, Laboratory of Livestock Genetics, Immunology and Physiology, Department Animal Production, Kasteelpark Arenberg 30, B-3001 Leuven, Belgium; email: Veerle.Bruggeman@biw.kuleuven.be C. Butter, Division of Immunology, Institute for Animal Health, Compton, N.A. Ciccone, Department of Veterinary Basic Sciences, The Royal Veterinary College, Camden, London NW1 0TU M. Clinton, The Roslin Institute and Royal (Dick) School of Veterinary mike.clinton@bbsrc. ac.uk S.R. Collett, The University of Georgia, College of Veterinary Medicine, T. Cserep, Intervet UK, Walton Manor, Walton, Milton Keynes, MK7 7AJ, UK; email: tibor.cserep@sp.intervet.com E. Decuypere, K.U.Leuven, Faculty of Bioscience Engineering, Depart ment of Biosystems, Laboratory of Livestock Genetics, Immunology and Physiology, Department Animal Production, Kasteelpark Arenberg 30, B-3001 Leuven, Belgium; email: eddy.decuypere@biw.kuleuven.be D.C. Deeming, Department of Biological Sciences, University of Lincoln, cdeeming@lincoln.ac.uk I.J.H. Duncan, Department of Animal and Poultry Science, University of iduncan@uoguelph. ca I.C. Dunn, Department of Genetics and Genomics, The Roslin Institute and 9PS, UK; email: ian.dunn@roslin.ed.ac.uk Contributors ix I. Estevez, Neiker-Tecnalia, Arkaute's Agrifood Campus, PO Box 46, C. Fisher, N.A. French, Aviagen Turkeys Ltd, Chowley Five, Chowley Oak Business Park, Tattenhall, Cheshire, CH3 9GA, UK; email: nfrench@aviagen.com R.T. Gladwell, School of Biological Sciences, The University of Reading, Whiteknights, Reading, RG6 6UB, UK; email: r.t.gladwell@reading. ac.uk R.M. Gous, University of KwaZulu-Natal, Pietermaritzburg, South Africa; email: gous@ukzn.ac.za P.M. Hocking, Division of Genetics and Genomics, The Roslin Institute and 9PS, UK; email: paul.hocking@roslin.ed.ac.uk N.T. Joseph, Department of Genetics and Genomics, The Roslin Institute 9PS, UK; email: nerine.joseph@roslin.ed.ac.uk M.T. Kidd, Mississippi State University, Department of Poultry Science, msstate.edu P.G. Knight, School of Biological Sciences, The University of Reading, Whiteknights, Reading, RG6 6UB, UK; email: p.g.knight@reading.ac.uk K.F. Laughlin, klaughlin@aviagen.com P.D. Lewis, Northcot, Cowdon Lane, Goodworth Clatford, Andover, SP11 7HG, UK; email: pdlewis@dsl.pipex.com T.M. Lovell, School of Biological Sciences, The University of Reading, Whiteknights, Reading, RG6 6UB, UK; email: t.m.lovell@reading.ac.uk J.C. McKay, UK; email: jim.mckay@ew-group.de S. Nandi, The Roslin Institute and Royal (Dick) School of Veterinary Studies, sunil.nandi@bbsrc.ac.uk O.M. Onagbesan, K.U.Leuven, Faculty of Bioscience Engineering, Depart- ment of Biosystems, Laboratory of Livestock Genetics, Immunology and Physiology, Department Animal Production, Kasteelpark Arenberg 30, B-3001 Leuven, Belgium; email: onagbesanok@yahoo.com T. Pizzari, Edward Grey Institute of Ornithology, Department of Zoology, tommaso.pizzari@zoology.oxford.ac.uk H.M. Sang, The Roslin Institute and Royal (Dick) School of Veterinary helen.sang@roslin. ed.ac.uk P.J. Sharp, Department of Genetics and Genomics, The Roslin Institute and x Contributors Royal (Dick) School of Veterinary Studies, University of Edinburgh, K. Tona, University of Lomé, Faculty of Sciences, Department of Animal H.J. Walter, Division of Immunology, Institute for Animal Health, Compton, com G.J. Wishart, Division of Biotechnology, University of Abertay, Bell Street, Dundee, BB1 1HG, UK [...]... been the foundation of a modern poultry industry that is a major source of animal protein in most countries of the world The history of poultry domestication and the development of a modern poultry industry are well reviewed (Crawford, 1990) Recent developments in knowledge and technology have changed the dynamics of poultry breeding © CAB International 2009 Biology of Breeding Poultry (ed P.M Hocking)... affected the reproductive systems of females and the mating efficiency of males The management and husbandry systems for breeding birds have also developed in parallel with the genetic changes, and a review of the current scientific knowledge of these birds is both timely and opportune This book contains reviews of the literature pertaining to breeding poultry of the three main poultry species (broiler, turkey... standards THE MODERN POULTRY INDUSTRY The production of poultry meat and eggs is a worldwide industry which supplies at least one-third of the animal-derived food for the 6 billion people on earth The statistical service of the Food and Agriculture Organization records that in 1961 the world produced less than 10 million tonnes of poultry meat By 2006 the world’s production of poultry meat was 81 million... Genetics of Modern Commercial Poultry 7 Continued improvements in poultry should be faster than in other species because poultry breeders have advantages of large population size, short generation interval and considerable genetic variation available to them THE FUTURE OF GENETICS IN COMMERCIAL POULTRY Breeding companies have the responsibility to manage their genetic resources to deliver stock of predictable... available 30 years ago shows that in 1975 it took 2.4 t of feed to produce each tonne of eggs whereas today it takes 1.9 t of feed to produce 1 t of eggs (Hy-Line and FAO: http://faostat fao.org) Today at least 115 million t of feed is used to produce eggs Using the 1975 genotypes to produce all of today’s eggs would require 144 million t of feed, an increase of 26% The genetic improvements in efficiency are... growing proportion of the world’s population A recent study in Australia has examined the sustainability of animal production industries in light of the growing concern about the environmental impact of various production systems (Foran et al., 2005) By taking account of all inputs and outputs they compare the greenhouse gas emissions of beef, lamb and pork production with that of poultry meat and eggs... production 3 Poultry health The development of effective vaccines and therapeutics, improved biosecurity and better nutrition have all contributed to improved health The emergence of breeding companies which are able to supply stock reliably free of the major vertically transmitted pathogens means that replacement stock can always be of a high health status Genetics of Modern Commercial Poultry 5 4... fertility and incubation The rest of the book covers the management of breeding birds: lighting and environmental enrichment (Part VI), nutrition (Part VII) and health (Part VIII) xi xii Preface The symposium, the 29th in the Poultry Science Symposium series, was held on 23–25 July 2007 at Surgeons’ Hall, Edinburgh, and consisted of short overviews of the material by each of the authors Unfortunately Dr... products of the industry available to a higher proportion of the world’s population The improvements in broiler efficiency are even more dramatic Between 1975 and today the combined effects of selection for growth, efficiency, yield and liveability have reduced the feed requirement for meat production from 20 million t of feed per million tonnes of meat to 8.5 million t of feed per million tonnes of meat... tonnes of meat (Aviagen and FAO) The genetic potential of birds is even better but is not realized in all production environments It took approximately 700 million t of feed to produce the 81 million t of poultry meat in 2005 Using a 1970s genotype would have required 1600 million t, an increase of 128% The annual improvement of 2–3% in efficiency of meat production has made a huge cumulative impact on . Poultry* 15 Meat Quality in Poultry and Game Birds 16 Avian Immunology 17 Reproductive Biology of Poultry 18 Poultry Genetics and Breeding 19 Nutrient Requirements of Poultry and Nutritional Research* 20. Bone Biology and Skeletal Disorders 24 Poultry Immunology* 25 Poultry Meat Science 26 Poultry Feedstuffs 27 Welfare of the Laying Hen 28 Avian Gut Function in Health and Disease 29 Biology of Breeding. foundation of a modern poultry industry that is a major source of animal protein in most countries of the world. The history of poultry domestication and the development of a modern poultry industry