Edited by Elaine Ostrander and Anatoly Ruvinsky The Genetics of the Dog, 2nd Edition FSC wwwracorg MIX Paper from responaible sources FSC CO13504 This page intentionally left blank The Genetics of the Dog, 2nd Edition Edited by Elaine A. Ostrander National Human Genome Research Institute National Institutes of Health Maryland USA and Anatoly Ruvinsky University of New England Australia 0 IY) www.cabi.org CABI is a trading name of CAB International CABI CABI Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491832111 Tel: +1 6173954056 Fax: +44 (0)1491833508 Fax: +1 6173546875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © CAB International 2012. 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 The genetics of the dog / edited by Elaine A. Ostrander and Anatoly Ruvinsky. -- 2nd ed. P. ;CM. Rev. ed. of: The genetics of the dog / edited by A. Ruvinsky and J. Sampson. c2001. Includes bibliographical references and index. ISBN 978-1-84593-940-3 (hardback : alk. paper) I. Ostrander, Elaine A. II. Ruvinsky, Anatoly. [DNLM: 1. Dogs--genetics. 2. Breeding. SF 427.2] LC classification not assigned 636.7'0821--dc23 2011031350 ISBN-13: 978 1 84593 940 3 Commissioning editor: Sarah Hulbert Editorial assistant: Gwenan Spearing Production editor: Fiona Chippendale Typeset by SPi, Pondicherry, India Printed and bound in the UK by CPI Group (UK) Ltd, Croydon, CR0 4YY I Contents Contributors Preface Elaine A. Ostrander and Anatoly Ruvinsky 1 Canid Phylogeny and Origin of the Domestic Dog 1 Caries Vila and Jennifer A. Leonard 2Experimental Studies of Early Canid Domestication 12 Lyudmila N. Trut, Irina N. Oskina and Anastasiya V Kharlamova 3 The History and Relationships of Dog Breeds 38 Heidi G. Parker 4 Molecular Genetics of Coat Colour, Texture and Length in the Dog 57 Christopher B. Kaelin and Gregory S. Barsh 5Mendelian Traits in the Dog 83 Frank W Nicholas 6 Canine Immunogenetics 91 Lorna J. Kennedy, William E.R. Oilier, Eliane Marti, John L. Wagner and Rainer E Storb 7 The Genetics of Canine Orthopaedic Traits 136 Gert J. Breur, Nicolaas E. Lambrechts and Rory J. Todhunter 8 Genetics of Cancer in Dogs 161 David R. Sargon 9Genetics of Neurological Disease in the Dog 189 Hannes Lohi 10 Genetics of Eye Disorders in the Dog 218 Cathryn S. Mellersh vi Contents) 11Canine Cytogenetics and Chromosome Maps 241 Matthew Breen and Rachael Thomas 12 Canine Genomics 255 Kerstin Lindblad-Toh 13Genetics of Canine Behavioural Disorders 275 Jennifer S. Yokoyama and Steven P Hamilton 14 Biology of Reproduction and Modern Reproductive Technology in the Dog 295 Catharina Linde Forsberg and Karine Reynaud 15 Developmental Genetics 321 Anatoly Ruvinsky and Mark Hill 16Genetics of Morphological Traits in the Domestic Dog 359 Elaine A. Ostrander and Carlos D. Bustamante 17 Canine Olfactory Genetics 375 Pascale Quignon, Stephanie Robin and Francis Galibert 18 Pedigree Analysis, Genotype Testing and Genetic Counselling 394 Anita M. Oberbauer 19Genetics of Quantitative Traits and Improvement of the Dog Breeds 421 Thomas R. Famula 20 Complex Traits in the Dog 435 Karl G. Lark and Kevin Chase 21 The Canine Model in Medical Genetics 458 Alan N. Wilton and Paula S. Henthorn 22 Genetic Aspects of Performance in Working Dogs 477 Heather J. Huson 23 Genetic Nomenclature 496 Zhi-Liang Hu and James M. Reecy Index 505 The colour plate section can be found following p. 244. I Contributors Gregory S. Barsh, Hudson Alpha Institute for Biotechnology, Huntsville, AL 35806 and Department of Genetics, Stanford University, Stanford, CA 94305, USA. Matthew Breen, Department of Molecular Biomedical Sciences, College of Veterinary Medicine and Center for Comparative Medicine and Translational Research, North Carolina State University, Raleigh, NC 27606, USA; also Lineberger Comprehensive Cancer Center, University of North Carolina, Chapel Hill, NC 27599, USA. Gert J. Breur, Department of Veterinary Clinical Sciences, School of Veterinary Medicine, Purdue University, West Lafayette, IN 47907, USA. Carlos D. Bustamante, Department of Genetics, Stanford University School of Medicine, Stanford, CA 94305, USA. Kevin Chase, Department of Biology, University of Utah, Salt Lake City, UT 84112, USA. Thomas R. Famula, Department of Animal Science, University of California Davis, CA 95616, USA. Francis Galibert, Institut de Genetique et Developpement de Rennes, UMR6061 CNRS Universite de Rennes 1, Faculte de Medecine, 2 avenue Prof. Leon Bernard, CS34317, 35043 Rennes Cedex, France. Steven P. Hamilton, Department of Psychiatry and Institute for Human Genetics, University of California, San Francisco, CA 94143-0984, USA. Paula S. Henthorn, Section of Medical Genetics, University of Pennsylvania School of Veterinary Medicine, Philadelphia, PA 19104-6010, USA. Mark Hill, Cell Biology Laboratory, School of Medical Sciences, University of New South Wales, Sydney, NSW, Australia. Zhi-Liang Hu, Department of Animal Science, Iowa State University, Ames, IA 50011, USA. Heather J. Huson, National Human Genome Research Institute, National Institutes of Health, 50 South Drive, Building 50, Room 5351, Bethesda, MD 20892 and Institute of Arctic Biology, University of Alaska, Fairbanks, AK 99775, USA. Christopher B. Kaelin, Hudson Alpha Institute for Biotechnology, Huntsville, AL 35806 and Department of Genetics, Stanford University, Stanford, CA 94305, USA. Lorna J. Kennedy, Centre for Integrated Genomic Medical Research, University of Manchester, Stopford Building Oxford Road, Manchester, M13 9PT, UK. Anastasiya V. Kharlamova, Institute of Cytology and Genetics, Siberian Branch of the Russian Academy of Sciences, 630090, Novosibirsk 630090, Russia. Nicolaas E. Lambrechts, Department of Veterinary Clinical Sciences, School of Veterinary Medicine, Purdue University, West Lafayette, IN 47907, USA. Karl G. Lark, Department of Biology, University of Utah, 257 South 1400 East, Salt Lake City, UT 84112, USA. vii viii Contributors) Jennifer A. Leonard, Conservation and Evolutionary Genetics Group, Estacien Biolegica de Doriana (EBD-CSIC), Avd. Americo Vespucio s/n, 41092 Seville, Spain. Kerstin Lindblad-Toh, Broad Institute of MIT and Harvard, Cambridge, MA 02142 and Science for Life Laboratory, Uppsala University, Uppsala Biomedicinska Centrum (BMC), Husargatan 3, 751 23 Uppsala, Sweden. Catharina Linde Forsberg, Department of Clinical Sciences, Division of Reproduction, Faculty of Veterinary Medicine and Animal Sciences, Swedish University of Agricultural Sciences, SE-750 07 Uppsala, Sweden. Hannes Lohi, Department of Veterinary Biosciences, Faculty of Veterinary Medicine and Research Programs Unit, Molecular Medicine, Faculty of Medicine, University of Helsinki; also Folkhalsan Research Center, Helsinki. Biomedicum 1, Room C323b, P.O. Box 63 (Haartmaninkatu 8), FI-00014 University of Helsinki, Finland. Eliane I. Marti, Universitat Bern, Abteilung Experimentelle Klinische Forschung, Langgassstrasse 124, 3001 Bern, Switzerland. Cathryn S. Mellersh, Animal Health Trust, Lanwades Park, Kentford, Newmarket, Suffolk, CB8 7UU, UK. Frank W. Nicholas, Faculty of Veterinary Science, University of Sydney, NSW 2006, Australia. Anita M. Oberbauer, Department of Animal Science, University of California Davis, CA 95616, USA. William E.R. Oilier, Centre for Integrated Genomic Medical Research, University of Manchester, Stopford Building, Oxford Road, Manchester, M13 9PT, UK. Irina N. Oskina, Institute of Cytology and Genetics, Siberian Branch of the Russian Academy of Sciences, 630090, Novosibirsk 630090, Russia. Elaine A. Ostrander, Cancer Genetics Section, National Human Genome Research Institute, National Institutes of Health, 50 South Drive, Building 50, Room 5351, Bethesda, MD 20892, USA. Heidi G. Parker, National Human Genome Research Institute, National Institutes of Health, 50 South Drive, Building 50, Room 5351, Bethesda, MD 20892, USA. Pascale Quignon, Institut de Genetique et Developpement de Rennes, UMR6061 CNRS Universite de Rennes 1, Faculte de Medecine, 2 avenue Prof. Leon Bernard, CS34317, 35043 Rennes Cedex, France. James M. Reecy, Department of Animal Science, Iowa State University, Ames, IA 50011, USA. Karine Reynaud, Reproduction and Developmental Biology, UMR 1198 INRA-ENVA, National Institute for Agricultural Research/National Veterinary School of Alfort, 7 Avenue General de Gaulle, 94700 Maisons-Alfort, France. Stephanie Robin, Institut de Genetique et Developpement de Rennes, UMR6061 CNRS Universite de Rennes 1, Faculte de Medecine, 2 avenue Prof. Leon Bernard, CS34317, 35043 Rennes Cedex, France. Anatoly Ruvinsky, University of New England, Armidale, 2351 NSW, Australia. David R. Sargan, Department of Veterinary Medicine, University of Cambridge, Madingley Road, Cambridge, CB3 OES, UK. Rainer E Storb, Transplantation Biology, Clinical Research Division, Fred Hutchinson Cancer Research Center, andf University of Washington School of Medicine, Department of Medicine, 1100 Fairview Ave. N., Seattle, WA 98109, USA. Rachael Thomas, Department of Molecular Biomedical Sciences, College of Veterinary Medicine and Center for Comparative Medicine and Translational Research, North Carolina State University, Raleigh,