Contributors Keith J. Betteridge Robert Gilmore McKinnell Benjamin G. Brackett Clement L. Markert Wallis Η. Clark, Jr. J. B. Peters Robert H. Foote George E. Seidel, Jr. W. C. D. Hare Sarah M. Seidel E. Keith Inskeep Thomas J. Sexton S. P. Leibo Elizabeth L. Singh Ann B. McGuire Davor Solter L. Dale Van Vleck New Technologies in Animal Breeding Edited by BENJAMIN G. BRACKET! Department of Clinical Studies-New Bolton Center School of Veterinary Medicine, and Department of Obstetrics and Gynecology School of Medicine University of Pennsylvania Kennett Square, Pennsylvania GEORGE E. SEIDEL, JR. Animal Reproduction Laboratory Colorado State University Fort Collins, Colorado SARAH M. SEIDEL Animal Reproduction Laboratory Colorado State University Fort Collins, Colorado 1981 ACADEMIC PRESS A Subsidiary of Harcourt Brace Jovanovich, Publishers NEW YORK LONDON PARIS SAN DIEGO SAN FRANCISCO SÄO PAULO SYDNEY TOKYO TORONTO This book was produced in part from work funded by the Office of Technology Assessment (OTA) of the United States Congress. The views expressed do not necessarily represent those of OTA. COPYRIGHT © 1981, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: New technologies in animal breeding. Includes bibliographies and index. 1. Livestock—Breeding. 2. Breeding. I. Brackett, Benjamin G. II. Seidel, George Ε., Jr. III. Seidel, Sarah Μ. IV. Title: Animal breeding. SF105.N484 636.082 81-20556 ISBN 0-12-123450-9 AACR2 PRINTED IN THE UNITED STATES OF AMERICA 81 82 83 84 9 8 7 6 5 4 3 2 1 Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. Keith J. Betteridge* (109), Agriculture Canada, Animal Diseases Research Insti tute, Nepean, Ontario K2H 8P9, Canada Benjamin G. Bracken (3, 141, 257), Department of Clinical Studies-New Bolton Center, School of Veterinary Medicine, and Department of Obstetrics and Gynecology, School of Medicine, University of Pennsylvania, Kennett Square, Pennsylvania 19348 Wallis Η. Clark, Jr. (91), Animal Science Department, University of California, Davis, California 95616 Robert H. Foote (13), Department of Animal Science, Cornell University, Ithaca, New York 14853 W. C. D. Hare (109), Agriculture Canada, Animal Diseases Research Institute - Nepean, Ontario K2H 89P, Canada E. Keith Inskeep (243), Division of Animal Science, West Virginia University, Morgantown, West Virginia 26506 S. P. Leibot (127), Biology Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee Ann B. McGuire (91), Animal Science Department, University of California, Davis, California 95616 Robert Gilmore McKinnell (163), Department of Genetics and Cell Biology, University of Minnesota, St. Paul, Minnesota 55108 * Present address: Centre de Recherche en Reproduction Animale, Faculte de Medecine Vet- erinaire, Universite de Montreal, C. P. 5000, St-Hyacinthe, Quebec J2S 7C6, Canada, tPresent address: Rio Vista International, Box 242, Route 9, San Antonio, Texas 78227. xi xii Contributors Clement L. Marken (181), Department of Biology, Yale University, New Haven, Connecticut 06511 J. B. Peters* (243), Division of Animal Science, West Virginia University, Morgantown, West Virginia 26506 George E. Seidel, Jr. (3,41, 181, 257), Animal Reproduction Laboratory, Colo rado State University, Fort Collins, Colorado 80523 Sarah M. Seidel (41, 257), Animal Reproduction Laboratory, Colorado State University, Fort Collins, Colorado 80523 Thomas J. Sexton (81),Avian Physiology Laboratory, U. S. Department of Agriculture, SEA-AR, Beltsville, Maryland 20705 Elizabeth L. Singh (109), Agriculture Canada, Animal Diseases Research Insti tute, Nepean, Ontario, Canada K2H 89P Davor Solter (201), The Wistar Institute of Anatomy and Biology, Philadelphia, Pennsylvania 19104 L. Dale Van Vleck (221), Department of Animal Science, Cornell University, Ithaca, New York 14853 * Deceased. Preface The desirability of reviewing new reproductive technologies in animal breed ing became a necessity upon initiation by the Office of Technology Assessment of the United States Congress (OTA) of a study of "Impacts of Applied Genetics: Animal Breeding" to enlighten legislative decision-makers. In the course of this assessment, 37 experts representing diverse areas convened in Denver, Colorado, January 15-17, 1980, for a conference from which this book has evolved. This volume is intended not to provide a comprehensive treatise of technologies of importance in breeding of domestic animals but to provide the reader with an appreciation for "what's new" within the framework of the animal breeding industry along with relevant implications. In early chapters, this volume brings together concise summations of the state-of-the art, along with thoughts on future directions, for several animal industries (artificial insemina tion, embryo transfer, poultry breeding, and aquaculture) of great importance in the production of food. In the middle chapters, developing technologies (sex selection, frozen storage of oocytes and embryos, in vitro fertilization and em bryo culture, amphibian nuclear transplantation, parthenogenesis, identical twins and cloning in mammals, and gene transfer in mammalian cells) along with potential applications and impacts on animal production are presented. In final chapters, analyses of potential genetic impacts (artificial insemination, sex selec tion, embryo transfer, cloning, and selfing in dairy cattle) and special economic considerations (benefits of reproductive management, synchronization of estrus, and artificial insemination of beef cattle and sheep) are presented, cases for which such illustrations were deemed most appropriate. The 14 chapters were written by 17 specialists representing a similar number of disciplines, but pulled together by a common interest in the intriguing possibilities for rapid extension of xiii xiv Preface basic knowledge generated through research to very practical applications in animal breeding. Emphasis is on food-producing and large domestic animals, especially the bovine species. Reproductive processes in these animals represent the most appropriate targets for application of the new technologies presented. As this book goes to press, feelings of sadness are shared with the family and many friends of Dr. J. B. Peters, a fellow contributor and esteemed colleague, recently deceased. Dr. Peters will be remembered for his devotion to the im provement of agriculture and for his good work in animal science at West Virginia University. The editors gratefully acknowledge the excellence and cooperation of each of the contributors, and the input of every participant in the OTA Conference on ^Impacts of Applied Genetics: Animal Breeding," and of scientists far and wide who have generously contributed information to the OTA assessment and to the literature—with apologies in advance for failure, in many instances, to make more proper acknowledgments. Special thanks are due to Miss Pamela J. Salsbury for secretarial assistance and to Mr. Lawrence Burton, Dr. Gretchen S. Kolsrud, and Dr. Zsolt Harsanyi of the Genetics and Population Program, OTA, U.S. Congress, for their effective collaboration in the early stages of this project. The interest, encouragement, and cooperation of the staff of Academic Press in expeditious publication are greatly appreciated. Benjamin G. Brackett George E. Seidel, Jr. Sarah M. Seidel 1 Perspectives on Animal Breeding GEORGE E. SEIDEL, JR., AND BENJAMIN G. BRACKETT I. Overview of Animals in Society 3 A. Role of Animals in Human Society 3 B. Competition between People and Animals for Food 4 C. Applications of Genetic Improvement 4 II. Perspectives on Animal Breeding 5 A. Interactions between Genotype and Environment 5 B. Characteristics of Genetic Improvement 5 C. Beginnings of Animal Breeding 6 D. Interdisciplinary Aspects of Animal Breeding 6 E. Problems of Animal Breeders 7 F. Circumventing Chance in Sexual Reproduction 7 G. Special Opportunities 8 III. Conclusions 9 References 9 I. OVERVIEW OF ANIMALS IN SOCIETY A. Role of Animals in Human Society HHoommoo ssaappiieennss hhaass eevvoollvveedd iinn aann eennvviirroonnmmeenntt wwiitthh ootthheerr aanniimmaallss.. HHee hhaass ddeeppeennddeedd oonn tthheemm ffoorr ffoooodd,, hhaaiirr aanndd hhiiddee,, ppoowweerr,, ssppoorrtt,, pprreessttiiggee,, aanndd ccoommppaann iioonnsshhiipp.. TThhee ddeeggrreeee ooff ddeeppeennddeennccee vvaarriieess aammoonngg ccuullttuurreess aanndd ggrroouuppss wwiitthhiinn ccuullttuurreess,, bbuutt tthhee ddeeppeennddeennccyy hhaass bbeeeenn aabbssoolluuttee iinn tteerrmmss ooff aatt lleeaasstt oonnee eesssseennttiiaall 33 NEW TECHNOLOGIES IN ANIMAL BREEDING Copyright © 1981 by Academic Press, Inc. All rights of reproduction in any form reserved. ISBN 0-12-123450-9 4 George Ε. Seidel, Jr., and Benjamin G. Brackett nutrient, vitamin B , which is not found in nature outside of animal tissues. 12 Furthermore, in the United States, animal products provide 69% of the protein in the average human diet, 80% of the calcium, about two-thirds of the phosphorus and vitamin B , the majority of vitamin B , as well as a substantial minority of 2 6 most other nutrients (Harper et al., 1980). Clearly, many human societies could survive without animals, but this would be both difficult and artificial, particu larly with nonaffluent cultures. Intangible values associated with man's relationships with animals, especially dogs and cats, often have profound emotional and psychological influences. Such influences have improved the quality of life for many, most strikingly for the very young, for the very old, and for those with certain kinds of mental illness. The following pages deal mainly with large, domestic animals, but food-producing species, for which the application of new technologies to en hance breeding might find encouragement from human society worldwide, will also be discussed. The important role of laboratory animals, e.g., frogs, hamsters, mice, rats, and rabbits, as experimental models to improve efficiency in animal breeding should be appreciated. The use of animals with appropriate genetic characteristics for studying human disease, nutrition, and other health- related problems is also of paramount importance to human society. B. Competition between People and Animals for Food It is often less efficient to feed plant products to animals than to consume them directly. On the other hand, the most abundant plant material, cellulose, is of little direct nutritive value to man, whereas cellulose is an excellent feed for ruminants and animals with a functional caecum. Domestic animals frequently eat what is aesthetically objectionable, of marginal or no food value, or even harmful to man, e.g., grass along the road, garbage, acorns, brewers' grains, beet pulp, citrus pulp, straw, cottonseed hulls, and stalk silage. It is true that animals compete with people for grain, but this argument has frequently been presented in a simplistic manner. A small amount of grain (or other relatively high energy substances, including some by-products) can greatly amplify the amount of food produced by animals that eat primarily nongrain feed. Fur thermore, when grain becomes scarce and prices increase, much less is fed to animals. This happened with dairy cattle in 1972 (see Fig. 3 in Chapter 2, this volume). Animals, therefore, act as a buffer for grain prices and supplies. When there are surpluses, animals utilize the excess. When there is scarcity, alternate feed supplies are used and/or animal numbers are reduced. C. Applications of Genetic Improvement One can logically conclude that domestic animals are likely to continue to be essential in most human societies and that their continued genetic improvement is 1. Perspectives on Animal Breeding 5 worthwhile. By and large, this book is limited to considerations of how to apply new technologies to food-producing domestic animals. However, many of the technologies will also be applicable to companion animals, circus and zoo ani mals, laboratory animals used for agricultural, biomedical, and industrial re search, and game and nongame wild animals (including avian and aquatic species). The technologies described may be particularly effective in rescuing some species from extinction or even possibly in recreating extinct species. The technologies also can contribute to greater quantities and improved quality of animal by-products, which frequently are worth more than the food component, e.g., wool, hides, aggressive behavior at rodeos, and velvet from antlers for use as an aphrodisiac in the Orient. II. PERSPECTIVES ON ANIMAL BREEDING A. Interactions between Genotype and Environment Why improve, or at least modify, the genetic makeup of animals? Fortunately, the classical arguments of genotype versus environment are obsolete. It is gener ally recognized that the environment is extremely important in producing a phenotypic product, such as milk or meat. However, in order to obtain a desired phenotype, both the genotype and the environment must be considered jointly. For example, poultry, regardless of genetic makeup, would freeze to death in winter in much of North America if not protected. Similarly, while dairy cows have calves that can be used for meat, such cows do extremely poorly on the western range in North America because they cannot find enough feed to have normal reproductive cycles and provide milk for the calf too; the genetic propen sity to divert nutrients to milk results in a huge nursing calf but no pregnancy for the next year. Conversely, although beef cows produce milk, they would quickly be culled in a dairy herd because of being unprofitable, i.e., they are not geneti cally geared to turn the large quantities of nutrients into large quantities of milk. It should be noted, however, that, although interactions between genotype and environment are important, they are not so exacting that strains developed in one environment cannot be used to advantage in some others. For example, bulls whose progeny are ranked differently in terms of milk production maintain the same rankings regardless of whether the cows are kept in North America, central Europe, southern Europe, Israel, or subtropical Latin America (see McDowell etal., 1976.) B. Characteristics of Genetic Improvement Although it is important to provide optimal environments for animal produc tion, animal breeding is aimed at providing optimal genotypes for the available