Biology of Breeding Poultry potx

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 Dig

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

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, UK

CABI is a trading name of CAB International

CABI Head Office

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.

Prospects for Sex Determination in Poultry 54

S Nandi and M Clinton

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

CHAPTER 8

Mating Behaviour and Fertility 111

I.J.H Duncan

CHAPTER 9

Sperm Competition and Fertilization Success 133

T.R Birkhead and T Pizzari

Hatching Egg and Chick Quality 224

V Bruggeman, K Tona, O Onagbesan and E Decuypere

P ART VI Managing the Environment 241

CHAPTER 14

Photoperiod and Control of Breeding 243

P.D Lewis

Protein and Amino Acid Responses 331

C Fisher and R.M Gous

Immune Protection of the Hatchling 391

C Butter and H.J Walter

C ONTRIBUTORS

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

Shefield, Shefield, S10 2TN, UK; email T.R.Birkhead@shefield.ac.uk

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,

Newbury, Berks, RG20 7NN, UK; email: Colin.Butter@bbsrc.ac.uk

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

Studies, Roslin, Midlothian, EH25 9PS, UK; email: mike.clinton@bbsrc ac.uk

S.R Collett, The University of Georgia, College of Veterinary Medicine,

Poultry Diagnostic and Research Centre, 953 College Station Road, Athens, Georgia, 30602-4875, USA; email: colletts@uga.edu

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,

Riseholme Park, Lincoln, LN2 2LG, UK; email: cdeeming@lincoln.ac.uk

I.J.H Duncan, Department of Animal and Poultry Science, University of

Guelph, Guelph, Ontario, Canada N1G 2W1; email: iduncan@uoguelph ca

I.C Dunn, Department of Genetics and Genomics, The Roslin Institute and

Royal (Dick) School of Veterinary Studies, Roslin, Midlothian, EH25 9PS, UK; email: ian.dunn@roslin.ed.ac.uk

Contributors ix

I Estevez, Neiker-Tecnalia, Arkaute's Agrifood Campus, PO Box 46,

E-01080 Vitoria-Gasteiz, Araba, Spain; email: iestevez@neiker.net

C Fisher, 20 Longstaff Court, Hebden Bridge, West Yorkshire, HX7 6AB, UK; email: colinisher1@btconnect.com

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

Royal (Dick) School of Veterinary Studies, Roslin, Midlothian, EH25 9PS, UK; email: paul.hocking@roslin.ed.ac.uk

N.T Joseph, Department of Genetics and Genomics, The Roslin Institute

and Royal (Dick) School of Veterinary Studies, Roslin, Midlothian, EH25 9PS, UK; email: nerine.joseph@roslin.ed.ac.uk

M.T Kidd, Mississippi State University, Department of Poultry Science,

Mississippi State, Mississippi 39762, USA; email: mkidd@poultry 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, Aviagen, 11 Lochend Road, Newbridge, Midlothian, EH28 8SZ, UK; email: 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, EW Group, Lochend Road, Newbridge, Midlothian, EH28 9SZ, UK; email: jim.mckay@ew-group.de

S Nandi, The Roslin Institute and Royal (Dick) School of Veterinary Studies,

Roslin, Midlothian, EH25 9PS, UK; email: 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,

University of Oxford, South Parks Road, Oxford, OX1 3PS, UK; email: tommaso.pizzari@zoology.oxford.ac.uk

H.M Sang, The Roslin Institute and Royal (Dick) School of Veterinary

Sciences, Roslin, Midlothian, EH25 9PS, UK; email: 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, Roslin, Midlothian, EH25 9PS, UK; email: peter.sharp@bbsrc.ac.uk

K Tona, University of Lomé, Faculty of Sciences, Department of Animal

Physiology, BP 1515 Lomé, Togo

H.J Walter, Division of Immunology, Institute for Animal Health, Compton,

Newbury, Berks, RG20 7NN, UK; email: helenjskinner78@btinternet com

G.J Wishart, Division of Biotechnology, University of Abertay, Bell Street,

Dundee, BB1 1HG, UK

P REFACE

Commercial broilers, turkeys and ducks are largely the products of 50 years of organized genetic selection for growth, feed efficiency and carcass yields in North America and Western Europe This process has revolutionized the poultry industry and resulted in the efficient worldwide production of nutritious and healthy meat for the consumer The intensive production of poultry meat continues to expand in many parts of the world, particularly in the emerging economies of Brazil, China and India

Chicks, poults and ducklings necessarily require adult male and female birds that are also required to reproduce efficiently Adults of current meat breeding lines are so radically changed from traditional lines that gave rise to them that an essentially new class of farm livestock has been produced The high growth rates of these birds lead inevitably to high adult body weights, which have also 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 and duck) and a chapter on minor species for which there is some information (Chapter 16) Typically these birds are fed on cereal-based diets and are housed on deep litter with various standards of environmental control, depending on the climate and region The broiler chicken is probably more advanced genetically than any other species, and in general the husbandry and management of the other species are based on the broiler chicken model There is relatively little published information on the duck and even less on geese, both of which are kept in some countries with access to water for swimming and green plant material as

a feed Geese are not commonly kept in large intensive operations or indoors, and Romanov (1999) has reviewed the available literature Ducks are also kept for the production of foie gras in France, and further information is available in the report by Guemene and Guy (2004)

An overview of genetic selection and developments in the management of breeding birds is given in Part I, followed by three chapters in Part II that summarize current developments in genetic knowledge that may be useful in the future; Parts III, IV and V review current knowledge on reproduction, mating, 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)

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 John Kirby and Dr Rob Renema were unable to produce a manuscript for the book

I am greatly indebted to the organizing committee, who generously provided their expertise in the disparate fields encompassed by this book, including initial technical editing of the draft manuscripts I am grateful for the support and advice of John Parsons and Kelvin McCracken, respectively secretary and treasurer of the UK Branch of WPSA, and to Liz Archibald for her sterling administrative support in preparation for the symposium The organizing committee consisted of P.M Hocking (Chairman), J.A Parsons, K.J McCracken, J.A Ball, J.S Bentley, T.F Davison, K.J Laughlin, P.J Sharp and the late G.C Perry

Dr Graham Perry was regrettably taken terminally ill just before the symposium and Dr Peter Lake kindly chaired the session on Mating Behaviour and Fertility Dr Perry organized the two very successful preceding symposia and provided a great deal of helpful advice and encouragement in the planning

of this meeting His enthusiasm, commitment and cheerful personality will be greatly missed

Finally, I wish to thank the sponsors for their generous support for the symposium, without which it would not have been possible to meet and discourse over 3 days in the genial surroundings of Surgeons’ Hall

P.M Hocking

Edinburgh June 2008

REFERENCES

Guemene, D and Guy, G (2004) The past, present and future of force-feeding and ‘foie gras’

production Worlds Poultry Science Journal 60, 210–222.

Romanov, M.N (1999) Goose production efficiency as influenced by genotype, nutrition and

production systems Worlds Poultry Science Journal 55, 281–294.

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P ART I

Introduction

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© CAB International 2009 Biology of Breeding Poultry (ed P.M Hocking) 3

Egg production has been improved consistently since the late 1930s, and the industry continues to improve the efficiency of laying hen production by at least 1% per year This requires the simultaneous improvement of multiple traits, including egg number, egg size, liveability, persistency and mature body weight There is also continuing progress in uniformity of egg size and colour and freedom from defects In broilers, combined selection for growth, body com position, feed efficiency and liveability continues to deliver 2–3% improvement per year in the efficiency of meat production Other traits such

as robustness, specific and general disease resistance, and absence of metabolic defects have also contributed to this progress

INTRODUCTION

Poultry have been domesticated for thousands of years, and man has made many genetic changes during the process of domestication and since then by establishing local varieties and selecting for various traits The genetic progress made since the late 1950s has 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