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MANAGING GLOBAL GENETIC RESOURCES Livestock Committee on Managing Global Genetic Resources: Agricultural Imperatives Board on Agriculture National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1993

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NATIONAL ACADEMY PRESS 2101 Constitution Avenue, NW Washington, DC 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Insti- tute of Medicine. This material is based on work supported by the U.S. Department of Agriculture, Agricultural Research Service, under Agreement No. 59-32U4-6-75. Additional fund- ing was provided by Calgene, Inc.; Educational Foundation of America; Kellogg En- dowment Fund of the National Academy of Sciences and the Institute of Medicine; Monsanto Company; Pioneer Hi-Bred International, Inc.; Rockefeller Foundation; U.S. Agency for International Development; U.S. Forest Service; W. K. Kellogg Foundation; World Bank; and Basic Science Fund of the National Academy of Sciences, contribu- tors to which include the Atlantic Richfield Foundation, AT&T Bell Laboratories, BP America, Inc., Dow Chemical Company, E.I. duPont de Nemours & Company, IBM Corporation, Merck & Co., Inc., Monsanto Company, and Shell Oil Company Founda- tion. Dissemination activities were supported in part by the W. K. Kellogg Founda- tion. Library of Congress Cataloging-in-Publication Data Livestock / Committee on Managing Global Genetic Resources: Agricultural Imperatives, Board on Agriculture, National Research Council. p. cm.-(Managing global genetic resources) Includes bibliographical references (p. ) and index. ISBN 0-309-04394-8 1. Livestock-Germplasm resources. I. National Research Council (U.S.). Committee on Managing Global Genetic Resources: Agricultural Imperatives. II. Series. SF105.3.L58 1993 636.08'21-dc20 93-16715 CIP Copyright 1993 by the National Academy of Sciences. All rights reserved. Any opinions, findings, conclusions, or recommendations expressed in this publica- tion are those of the author(s) and do not necessarily reflect the view of the organiza- tions or agencies that provided support for this project. Printed in the United States of America

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Committee on Managing Global Genetic Resources: Agricultural Imperatives PETER R. DAY, Chair, Rutgers University ROBERT W. ALLARD, University of California, Davis PAULO DE T. ALVIM, Comissao Executiva do Plano da Lavoura Cacaueira, Brasil~ JOHN H. BARTON, Stanford University FREDERICK H. BUTTEL, University of Wisconsin TE-TZU CHANG, International Rice Research Institute, The Philippines (Retired) ROBERT E. EVENSON, Yale University HENRY A. FITZHUGH, International Livestock Center for Africa. Ethiopia MAJOR M. GOODMAN, North Carolina State University JAAP I. HARDON, Center for Genetic Resources, The Netherlands DONALD R. MARSHALL, University of Sydney, Australia SETIJATI SASTRAPRADJA, National Center for Biotechnology, Indonesia CHARLES SMITH, University of Guelph, Canada JOHN A. SPENCE, University of the West Indies, Trinidad and Tobago Genetic Resources Staff MICHAEL S. STRAUSS, Project Director JOHN A. PINO, Project Directort BRENDA BALLACHEY, Staff Of ficert BARBARA I. RICE, Project Associate and Editor Executive Commission of the Program for Strengthening Cacao Production, Brazil. "Through June 1990. "Through November 1989. . . . Itt

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Subcommittee on Animal Genetic Resources HENRY A. FITZHUGH, Chair, International Livestock Center for Africa, Ethiopia ELIZABETH L. HENSON, Cotswold Farm Park, England JOHN HODGES, Mittersill, Austria DAVID R. NOTTER, Virginia Polytechnic Institute and State University DIETER PLASSE, Universidad Central de Venezuela (Retired) LOUISE LETHOLA SETSHWAELO, Ministry of Agriculture, Botswana THOMAS E. WAGNER, Ohio University, Athens AWES E. WOMACK, Texas A&M University Rev

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Board on Agriculture THEODORE L. HULLAR, Chair, University of California, Davis PHILIP H. ABELSON, American Association for the Advancement of Science, Washington, D.C. JOHN M. ANTLE, Montana State University DALE E. BAUMAN, Cornell University WILLIAM B. DELAUDER, Delaware State College SUSAN K. HARLANDER, Land O'Lakes, Inc., Minneapolis, Minnesota PAUL W. JOHNSON, Natural Resources Consultant, Decorah, Iowa T. KENT KIRK, U.S. Department of Agriculture, Madison, Wisconsin JAMES R. MOSELEY, km Moseley Farms, Inc., Clark Hills, Indiana DONALD R. NIELSEN, University of California, Davis NORMAN R. SCOTT, Cornell University GEORGE E. SEIDEL, JR., Colorado State University PATRICIA B. SWAN, Iowa State University JOHN R. WELSERr The Upjohn Company, Kalamazoo, Michigan FREDERIC WINTHROP, JR., The Trustees of Reservations, Beverly, Massachusetts SUSAN E. OFFUTT, Executive Director JAMES E. TAVARES, Associate Executive Director CARLA CARLSON, Director of Communications BARBARA I. RICE, Editor JANET L. OVERTON, Associate Editor v

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The National Academy of Sciences is a private, nonprofit, self- perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the au- thority of the charter granted to it by the Congress in 1863, the Acad- emy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Frank Press is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its ad- ministration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the fed- eral government. The National Academy of Engineering also spon- sors engineering programs aimed at meeting national needs, encour- ages education and research, and recognizes the superior achievements of engineers. Dr. Robert M. White is president of the National Acad- emy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertain- ing to the health of the public. The Institute acts under the responsi- bility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Kenneth I. Shine is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and advising the federal government. Functioning in ac- cordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engi- neering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Frank Press and Dr. Robert M. White are chairman and vice-chairman, respectively, of the National Research Council. Al

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Preface Humans have tended sheep, pigs, cattle, chickens, and other domesticated animals as a source of food, tallow, leather, and draft power for more than 10,000 years. It was only as recently as the early twentieth century, however, that deliberate and planned selective breeding began to shift the genetic makeup of do- mesticated populations in predetermined directions. In some spe- cies, divergent goals were set for different populations. For example, cattle were selected and bred for meat or for milk production. In 1905, this division was recognized by the formation of the Dairy Shorthorn Society in Britain. The application of the science of genetics in this century to the breeding and improvement of livestock has yielded remarkable re- sults. For example, a single dairy cow in the developed world pro- duces nearly twice the amount of milk per year today than her ances- tor could produce just 25 years ago. Since 1983, in less than 10 years, pork producers have reduced fat in pigs by 31 percent and saturated fat by 29 percent. In fewer than 50 years, poultry producers in indus- trialized countries have reduced, by more than half, the number of days required to raise broiler chickens that are ready for market, from 95 days in 1934 to 45 days in 1991. Feeding and management improvements contribute to these per- formance achievements in food animals, but central to these develop- ments has been genetic improvement made possible by the availabil- ity of diverse breeds and populations that are the sources of genes needed for breed improvement. However, the focus in the agricul . . Ott

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viii / Preface tural sector has been toward fewer breeds, and breeds that are suited for wide distribution throughout the world. As these new breeds spread, they displace the many breeds and populations that were the genetic sources for their development. Improvements in animal breeding techniques and production have advanced rapidly. Perhaps even more rapidly during the past 10 years have come the developments in molecular biology and repro- ductive physiology that will eventually contribute to furler improvements in animal breeding and production and to the assurance of a quality food supply. Noting the swift advances in science and technology, the Committee on Managing Global Genetic Resources believes that now is the time to assess the goals and future directions of the live- stock research and production communities, with a specific focus on genetic diversity. Improvements in all areas of animal husbandry, including breeding and genetics, reproductive physiology, nutrition, and animal health, can unintentionally contribute to the decline in the number and diversity of breeds and types. Loss of genetic diver- sity threatens livestock production systems throughout the world. Preserving the capacity to continue to develop modern livestock re- quires global actions, on national and international levels, to prevent the loss of valuable genetic resources. During its assessment and deliberations, the committee observed that consensus exists within the livestock production community on the need to develop national and international efforts to conserve and manage livestock genetic resources. The approaches to cor~ser- vation, however, are markedly different and are generally defined by two philosophies: (1) utilization and (2) preservation. The utilizationist gives the highest priority to using available genetic resources to im- prove livestock populations. The objective is to increase the rate and efficiency of livestock food and fiber production. The preservationist emphasizes the value of preserving the widest possible spectrum of genetic diversity to meet future needs. Breed preservation is the preferred method of gene conservation. Reconciliation of these two views is needed, and it must begin with the recognition of their common goal sustained improvement of livestock for use by an expanding global population. The commit- tee believes that the information in Managing Global Genetic Resources: Livestock can provide the foundation for incorporating both views into a cohesive strategy for conservation and management. This report examines the genetic diversity of the major domestic livestock species. Those used on a global scale include cattle, sheep, goats, pigs, horses, donkeys, buffaloes, chickens, turkeys, ducks, and geese. The report assesses the status of genetic diversity in the most

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Preface / ix common of these livestock species and the need for actions to con- serve them as resources for future agricultural and livestock develop- ment. Although the report focuses on only a few animal species, it notes that many others, for example, camels, llamas, rabbits, and guinea pigs, are important in various countries or regions. In general, the prin- ciples and technologies of genetic conservation presented in this re- port are applicable to all agriculturally important domestic animals. The committee, established by the National Research Council's Board on Agriculture in November 1986, is concerned with genetic resources of identified economic value. These resources are impor- tant to agriculture, forestry, and industry. The committee has been assisted by two subcommittees and several work groups that gath- ered information or prepared specific reports. One of the subcom- mittees, chaired by Henry A. Fitzhugh of the International Livestock Center for Africa in Ethiopia, examined the conservation and man- agement of livestock genetic resources and drafted this report. It is one in a series of reports entitled, Managing Global Genetic Resources. Previously published reports in the series are The U.S. National Plant Germplasm System and Forest Trees, both published in 1991. The com- mittee believes that the status of the genetic diversity of commercial fish and shellfish is an important area of study, but the lack of re- search and documentation in this area precluded it from publishing a separate report. Soon to be released is the committee's final report, Agricultural Crop Issues and Policies. In addition to the examination of crop plants, the committee's main report will address the legal, political, economic, and social issues surrounding global genetic resources management as they relate to agricultural imperatives. The Subcommittee on Animal Genetic Resources was formed to assess policies and programs for preserving and using animal genetic resources and to identify strategies for meeting present and future needs. Specifically the subcommittee was asked to do the following: belly. Examine the uses and status of livestock genetic resources glo Examine methods for using and preserving livestock genetic resources. Identify the major problems that limit effective management of global resources. Assess the status of international efforts to conserve and ex- change animal germplasm. Recommend research and development priorities and practical

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x / Preface strategies for animal genetic resources management at national, re- gional, and global levels. Present a global strategy for conserving and managing live- stock genetic resources. Chapter 1 of this report discusses the need to conserve livestock genetic resources, especially in developing countries where many livestock populations have not been sufficiently characterized. It summarizes the utilizationist and preservationist approaches to conservation and emphasizes a position that would be acceptable to these divergent views. Chapter 2 describes the fundamental elements in livestock genetic resource management, including rationale for preserving popu- lations, identifying sampling strategies, and developing preservation methods. Chapter 3 defines the factors that affect genetic variation, and Chapter 4 profiles the biotechnological methods for delineating and manipulating genetic material. Chapter 5 describes examples and organizational elements of national programs that are involved with managing and conserving livestock genetic resources. Chapter 6 presents the committee's recommendations on strengthening inter- national programs for conservation of animal germplasm. Appendixes include an overview of the major livestock species by total population sizes and global distributions; a discussion of the risks of disease transmission during embryo transfer; and techniques for the collection, handling, and storage of semen. The issues surrounding the conservation animal genetic resources have been discussed and debated for more than 30 years. Conclu- sions generally have been similar diversity in livestock populations is at risk, valid reasons exist to conserve diversity, and organized conservation activities are highly desirable. Nevertheless, active pro- grams to inventory and conserve genetic diversity have been the ex- ception rather than the rule. Livestock genetic diversity is essential to meet the future needs of global society. Technologies that emerge from scientific research will enhance capacities to conserve and use animal genetic diversity. Ultimately, conservation of livestock genetic resources will require cooperation at the national and global levels. The committee be- lieves that the conclusions and recommendations presented in this report can contribute to improved conservation and management of a major global resource livestock. PETER R. DAY, Chair Committee on Managing Global Genetic Resources: Agricultural Imperatives

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Acknowledgments Many scientists and policymakers have contributed time, sup- port, and information instrumental to the committee's analyses con- tained in this report. In particular the committee thanks Keith Gre- gory, Gordon Dickerson, John Acree, Roger Gerrits, and Henry Shands for their advice and contributions to the study. Others have pro- vided valuable information and insight. These include Eric Bradford, Harvey D. Blackburn, ferry Callis, Thomas Cartwright, Roy Crawford, Dale Gustafson, William Hohenboken, Katie Maijala, Ian Mason, Charles Mullenax, Ian Parsonson, Caird Rexroad, Elizabeth Singh, Philip Sponenberg, lane Wheeler, and Saul Wilson. Many other scientists and policymakers assisted by reviewing selected draft materials and the appendixes. To all of these individuals the committee expresses its gratitude. Administrative support during various stages of the development of this report was provided by Philomina Mammen, Carole Spalding, Maryann Tully, Joseph Gagnier, and Mary Lou Sutton, and they are gratefully acknowledged. The committee also thanks Joi Brooks and Sherry Showell, interns sponsored by the Midwestern University Con- sortium for International Activities, for assisting in the development of the report. x~

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Contents EXECUTIVE SUMMARY Two Views on Livestock Conservation, 2 Genetic Diversity of Livestock, 3 Livestock Production Systems, 6 Why Conserve the Genetic Diversity of Livestock?, 9 Technologies for Conserving and Using Germplasm, 10 Methods of Preserving Animal Germplasm, 12 Efforts to Implement Livestock Conservation, 14 Recommendations, 15 1 1 THE NEED TO CONSERVE LIVESTOCK GENETIC RESOURCES 21 The Development of Animal Agriculture, 21 The Effects of Technology on Genetic Diversity, 26 Rationales for Conserving Livestock Genetic Resources, 38 Approaches to Livestock Conservation, 42 Recommendations, 46 2 ESSENTIAL CONSERVATION CONSIDERATIONS Criteria for Conserving Populations, 49 Strategies for Sampling Populations, 53 Methods for Preserving Livestock Germplasm, 54 Recommendations, 59 ....... 49 3 MEASUREMENT AND USE OF GENETIC VARIATION.... 63 The Influence of Human Society, 63 Factors Affecting Genetic Variation, 67 Germplasm Use, 69 Recommendations, 74 . . . x''`

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xiv / Contents 4 NEW TECHNOLOGY AND ITS IMPACT ON CONSERVATION ......... Methods to Quantify Genetic Variation, 77 ~T A T 1 ~ ~ w~^ curares ana Gene transfer Methodologies, 83 Reproductive Technologies, 86 Health Status of Germplasm, 94 Recommendations, 95 5 N A TIO N AL PR O G R. A M S 97 Examples of Current National Efforts, 97 Organizational Elements of a National Program, 101 Germplasm Conservation in Developed and Developing Countries, 106 Recommendations, 108 6 INTERNATIONAL PROGRAMS AND A GLOBAL MECHANISM................................ International Programs, 112 Information Sources and Data Bases, 118 Creating Regional Stores of Frozen Germplasm, 121 Related Issues of Plant Genetic Resource Programs, 122 Global Conservation of Animal Germplasm, 123 Recommendations, 128 REFERENCES ................... APPENDIXES A Global Status of Livestock and Poultry Species, 141 Ian L. Mason and Roy D. Crawford B Embryo Transfer: An Assessment of the Risks of Disease Transmission, 171 Elizabeth Singh Animal Genetic Resources: Sperm, 215 Ian Parsonson .. 111 ................ 131 GLOSSARY . ~245 ABBREVIATIONS 253 AUTHORS . . . INDEX

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Livestock

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