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Resource Allocation Theory Applied to Farm Animal Production

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Resource Allocation Theory Applied to Farm Animal Production RESOURCE ALLOCATION T HEORY APPLIED TO FARM ANIMAL PRODUCTION It is the magician’s wand, by means of which [the agriculturist] may summon into life whatever form and mould he pleases William Youatt (1872) cited in Darwin (1872) One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal’s ( . ) own good, but to man’s use or fancy Charles Darwin (1872) RESOURCE ALLOCATION THEORY APPLIED TO FARM ANIMAL PRODUCTION Edited by Wendy Mercedes Rauw Department of Animal Biotechnology University of Nevada-Reno Reno, USA CABI is a trading name of CAB International CABI Head Offi ce CABI North American Offi ce Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org ©CAB International 2009. All rights reserved. No part of this publication may be re- produced 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 Resource allocation theory applied to farm animal production / edited by Wendy M. Rauw. p. cm. Includes bibliographical references and index. ISBN 978-1-84593-394-4 (alk. paper) 1. Livestock productivity--Mathematical models. 2. Resource allocation. I. Rauw, Wendy M. II. Title. SF140.M35R47 2008 636'.08--dc22 2008025072 ISBN: 978 1 84593 394 4 Typeset by SPi, Pondicherry, India. Printed and bound in the UK by MPG Books Ltd, Bodmin. The paper used for the text pages in this book is FSC certifi ed. The FSC (Forest Stewardship Council) is an international network to promote responsible management of the world’s forests. Contents Contributors vii PART I RESOURCE ALLOCATION 1 Introduction 1 W.M. Rauw 2 Resource Allocation Patterns 22 D.S. Glazier 3 Trade-offs 44 D.S. Glazier 4 Metabolic Constraints to Resource Allocation 61 D.E. Naya and L.D. Bacigalupe 5 Homeorhesis During Heat Stress 72 R.J. Collier, S.W. Limesand, M.L. Rhoads, R.P. Rhoads and L.H. Baumgard PART II INPUTS AND OUTPUTS 6 Residual Feed Intake 89 R.M. Herd 7 Allocation of Resources to Maintenance 110 P.W. Knap 8 Allocation of Resources to Growth 130 C.T. Whittemore 9 Genetic Size-scaling 147 St C.S. Taylor v vi Contents 10 Allocation of Resources to Reproduction 169 G.B. Martin, D. Blache and I.H. Williams 11 Allocation of Resources to Immune Responses 192 I.G. Colditz PART III CONSEQUENCES OF SELECTION FOR INCREASED PRODUCTION EFFICIENCY 12 Selection for High Production in Pigs 210 P.W. Knap and W.M. Rauw 13 Selection for High Production in Poultry 230 P.B. Siegel, C.F. Honaker and W.M. Rauw 14 Selection for High Production in Dairy Cattle 243 R.F. Veerkamp, J.J. Windig, M.P.L. Calus, W. Ouweltjes, Y. de Haas and B. Beerda 15 Consequences of Biological Engineering 261 for Resource Allocation and Welfare D.M. Broom PART IV ANIMAL BREEDING AND RESOURCE MODELLING 16 Breeding Goals to Optimize Production Effi ciency 275 A.F. Groen 17 Robustness 288 P.W. Knap 18 Modelling of Resource Allocation Patterns 302 N.C. Friggens and E.H. van der Waaij Index 321 Contributors L.D. Bacigalupe, Department of Animal and Plant Sciences, University of Sheffi eld, Western Bank, Sheffi eld S10 2TN, UK. Present address: Departamento de Zoología, Universidad de Concepción, Casilla 160-C Concepción, Chile. L.H. Baumgard, Department of Animal Sciences, University of Arizona, Tucson, AZ 85721, USA. B. Beerda, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. D. Blache, UWA Institute of Agriculture M082, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia. D.M. Broom, Centre for Animal Welfare and Anthrozoology, Department of Veterinary Medicine, University of Cambridge, Madingley Road, Cambridge CB3 0ES, UK. M.P.L. Calus, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. I.G. Colditz, CSIRO Livestock Industries, FD McMaster Laboratory, Armidale, NSW 2350, Australia. R.J. Collier, Department of Animal Sciences, University of Arizona, Tucson, AZ 85721, USA. Y. de Haas, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. N.C. Friggens, Department of Animal Health, Welfare and Nutrition, Research Centre Fou- lum, 8830 Tjele, Denmark. D.S. Glazier, Department of Biology, Brumbaugh Academic Center, Juniata College, Huntingdon, PA 16652, USA. A.F. Groen, Director Corporate Staff Education and Research, Wageningen University and Research Center, PO Box 9101, 6700 HB Wageningen, The Netherlands. R.M. Herd, New South Wales Department of Primary Industries, Beef Industry Centre, Armidale, NSW 2351, Australia. C.F. Honaker, Department of Animal and Poultry Sciences, Virginia Tech, Blacksburg, VA 24061, USA. vii viii Contributors P.W. Knap, PIC International Group, 24837 Schleswig, Germany. S.W. Limesand, Department of Animal Sciences, University of Arizona, Tucson, AZ 85721, USA. G.B. Martin, UWA Institute of Agriculture M082, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia. D.E. Naya, Center for Advanced Studies in Ecology and Biodiversity and Departamento de Ecología, Facultad de Ciencias Biológicas, Pontifi cia Universidad Católica de Chile, CP 6513677, Santiago, Chile. W. Ouweltjes, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. W.M. Rauw, University of Nevada-Reno Department of Animal Biotechnology, Reno, NV 89557, USA. M.L. Rhoads, Department of Animal Sciences, University of Arizona, Tucson, AZ 85721, USA. R.P. Rhoads, Department of Animal Sciences, University of Arizona, Tucson, AZ 85721, USA. P.B. Siegel, Department of Animal and Poultry Sciences, Virginia Tech, Blacksburg, VA 24061, USA. St C.S. Taylor, Animal Breeding Research Organisation, Edinburgh, UK. E.H. van der Waaij, Wageningen University, Animal Breeding and Genomics Centre, 6700 AH W ageningen, The Netherlands. R.F. Veerkamp, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. C.T. Whittemore, University of Edinburgh, Edinburth EH9 3JG, UK. J. J. Windig, Animal Breeding and Genomics Centre, Wageningen University and Research Centre, 8200 AB Lelystad, The Netherlands. I.H. Williams, UWA Institute of Agriculture M082, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia. 1 Introduction W.M. RAUW University of Nevada-Reno, Department of Animal Biotechnology, Reno, NV 89557, USA 1. Life History Theory When Charles Darwin travelled with the H.M.S. Beagle to Tierra del Fuego and the Falkland Islands in 1834, he observed a discrepancy between the number of eggs of a large white Doris (a sea slug) and the abundance of the species. He real- ized that the abundance of a species does not necessarily depend on the number of offspring produced (Van Straalen and Roelofs, 2006). Life history theory deals with the question of why the power of propagation differs so much between spe- cies, and with the way an organism spreads its reproduction over its lifetime and forms an adaptation to the environment it lives in (Brommer, 2000; Van Straalen and Roelofs, 2006). Darwin adopted Herbert Spencer’s (1864) term ‘survival of the fittest’ in 1866. His concept of fitness arose from his view of the organism and the environment it lives in. He did not actually use the term ‘fitness’, but referred to individual organisms as being more or less ‘fit’ than other individuals: different individual members of a species ‘fit’ into the environment to different degrees as a consequence of phenotypic variation, and those that make the best ‘fit’ survive and reproduce their kind better than those whose ‘fit’ is poorer (Ariew and Lewontin, 2004). In other words, fit organisms are better represented in future generations than their relatively unfit competitors (Stearns, 1976). Textbooks, monographs and articles show a wealth of diversity in fitness definitions (De Jong, 1994). Or as Ariew and Lewontin (2004) state: ‘No concept in evolutionary biology has been more confusing and has produced such a rich philosophical literature as that of fitness.’ Fitness concepts may refer to the functioning of an organism (fitness itself is a cause of natural selection), or may consti- tute a technical term in population biology summarizing numerical processes (fitness is a description of natural selection; De Jong, 1994). Within the numerical fitness con- cept, many quantities are proposed as fitness measures: life-history traits may include lifetime reproductive success, survival, viability, fecundity, mating success and age at maturity (Schluter et al., 1991; De Jong, 1994). Theories on the evolution of life history ©CAB International 2009. Resource Allocation Theory Applied to Farm Animal Production (ed. W.M. Rauw) 1 2 W.M. Rauw focus on the notion that natural selection results in maximal fitness and prunes away less-optimal life histories (Brommer, 2000). Life histories are shaped by the interaction of extrinsic factors, i.e. environmen- tal impacts on survival and reproduction, and intrinsic factors, i.e. trade-offs among life-history traits and lineage-specific constrains on the expression of genetic varia- tion (Stearns, 2000). Cole (1954) referred to the extrinsic factors when stating: ‘It is obvious that the ability of the ancestors of existing species to replace themselves has been sufficient to overcome all environmental exigencies which have been encoun- tered (. .) through physiological, morphological, and behavioural adaptations that enable offspring to be produced and to survive in sufficient numbers to insure the persistence of a species.’ Thus, in the absence of trade-offs, selection would drive all life-history traits to limits imposed by design and history, i.e.
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