Nutrient Requirements of Nonhuman Primates

Second Revised Edition, 2003

Committee on Animal Nutrition

Ad Hoc Committee on Nonhuman Primate Nutrition

Board on Agriculture and Natural Resources

Division on Earth and Life Studies

NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES

THE NATIONAL ACADEMIES PRESS
Washington, D.C.
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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Nutrient Requirements of Nonhuman Primates Second Revised Edition, 2003 Committee on Animal Nutrition Ad Hoc Committee on Nonhuman Primate Nutrition Board on Agriculture and Natural Resources Division on Earth and Life Studies NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES THE NATIONAL ACADEMIES PRESS Washington, D.C. www.nap.edu

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 THE NATIONAL ACADEMIES PRESS 500 Fifth St., NW Washington, D.C. 20001 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 study was supported by the Department of Health and Human Services, the American Zoo and Aquarium Association, The Geraldine R. Dodge Foundation, Association of Primate Veterinarians, Harlan Teklad, Purina Mills, Inc., and Zupreem. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. Library of Congress Cataloging-in-Publication Data Nutrient requirements of nonhuman primates.—2nd rev. ed. p. cm.—(Nutrient requirements of animals) Rev. ed. of: Nutrient requirements of nonhuman primates / Panel on Nonhuman Primate Nutrition, Subcommittee on Laboratory Animal Nutrition, Committee on Animal Nutrition, Board on Agriculture and Renewable Resources, National Research Council. 1978. Includes bibliographical references (p. ). ISBN 0-309-06989-0 (pbk.) 1. Primates—Feeding and feeds. 2. Primates—Nutrition—Requirements. 3. Primates as laboratory animals. I. National Research Council (U.S.). Committee on Animal Nutrition. II. National Research Council (U.S.). Panel on Nonhuman Primate Nutrition. Nutrient requirements of nonhuman primates. III. Series. SF407.P7 N88 2002 636.98—dc21 2002013021 Additional copies of this report are available from the National Academies Press, 500 Fifth St., NW, Box 285, Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet http://www.nap.edu. Copyright 2003 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 THE NATIONAL ACADEMIES Advisers to the Nation on Science, Engineering, and Medicine 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 authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce M. Alberts 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 administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf is president of the National Academy 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 pertaining to the health of the public. The Institute acts under the responsibility 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. Harvey V. Fineberg 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 accordance 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 engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce M. Alberts and Dr. Wm. A. Wulf are chair and vice chair, respectively, of the National Research Council www.national-academies.org

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 AD HOC COMMITTEE ON NONHUMAN PRIMATE NUTRITION DUANE E. ULLREY, Chair, Michigan State University, East Lansing, Michigan MARY E. ALLEN, National Zoological Park, Washington, D.C. LYNNE M. AUSMAN, Tufts University, Boston, Massachusetts NANCY L. CONKLIN-BRITTAIN, Harvard University, Cambridge, Massachusetts MARK S. EDWARDS, Zoological Society of San Diego, San Diego, California JOSEPH M. ERWIN, Diagnon Corporation/Bioqual, Inc., Rockville, Maryland School of Medicine, University of Maryland, Baltimore MICHAEL F. HOLICK, Boston University School of Medicine, Massachusetts DANIEL T. HOPKINS, Purina Mills, Inc., retired, St. Charles, Missouri SHERRY M. LEWIS, National Center for Toxicological Research, The Bionetics Corporation, Jefferson, Arkansas BO L. G. LONNERDAL, University of California-Davis, Davis, California LAWRENCE L. RUDEL, Wake Forest School of Medicine, Winston-Salem, North Carolina COMMITTEE ON ANIMAL NUTRITION GARY L. CROMWELL, Chair, University of Kentucky MARY E. ALLEN, National Zoological Park, Washington, D.C. MICHAEL L. GALYEAN, West Texas A&M University RONALD W. HARDY, University of Idaho BRIAN W. McBRIDE, University of Guelph, Ontario, Canada KEITH RINEHART, Perdue Farms, Inc., Salisbury, Maryland L. LEE SOUTHERN, Louisiana State University JERRY W. SPEARS, North Carolina State University DONALD R. TOPLIFF, Oklahoma State University WILLIAM P. WEISS, The Ohio State University Staff CHARLOTTE KIRK BAER, Program Director NORMAN GROSSBLATT, Editor STEPHANIE PADGHAM, Project Assistant

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 BOARD ON AGRICULTURE AND NATURAL RESOURCES HARLEY W. MOON, Chair, Iowa State University CORNELIA B. FLORA, Iowa State University ROBERT B. FRIDLEY, University of California BARBARA GLENN, Federation of Animal Science Societies LINDA GOLODNER, National Consumers League W.R. (REG) GOMES, University of California PERRY R. HAGENSTEIN, Institute for Forest Analysis, Planning, and Policy, Wayland, Massachusetts GEORGE R. HALLBERG, The Cadmus Group, Inc. CALESTOUS JUMA, Harvard University GILBERT A. LEVEILLE, McNeil Consumer Healthcare, Denville, New Jersey WHITNEY MACMILLAN, Cargill, Inc., Minneapolis, Minnesota TERRY MEDLEY, DuPont Biosolutions Enterprise WILLIAM L. OGREN, U.S. Department of Agriculture ALICE PELL, Cornell University NANCY J. RACHMAN, Novigen Sciences, Inc. G. EDWARD SCHUH, University of Minnesota BRIAN STASKAWICZ, University of California, Berkeley JOHN W. SUTTIE, University of Wisconsin JAMES TUMLINSON, USDA, ARS JAMES J. ZUICHES, Washington State University Staff CHARLOTTE KIRK BAER, Director STEPHANIE PADGHAM, Administrative Assistant SHIRLEY B. THATCHER, Administrative Assistant* *   through March 2000

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Preface This report is one of a series issued under the direction of the National Research Council’s Committee on Animal Nutrition (CAN) of The National Academies Board on Agriculture and Natural Resources. It was prepared by the CAN Ad Hoc Committee on Nonhuman Primate Nutrition and is a revision of the 1978 edition of Nutrient Requirements of Nonhuman Primates. Throughout the study process, input from others has been sought by posing specific questions in widely distributed letters, by hosting workshops and information-gathering sessions, and by inviting sponsors and the general public to attend meetings of the Committee. Information published before 1978 has been reevaluated, that in newer publications has been examined, and both have been used to update this report. Greater emphasis than before has been placed on descriptions of natural dietary habits, gastrointestinal anatomy and physiology, and the special nutrient and dietary husbandry needs of species that traditionally have been difficult to maintain in captivity. The order Primates is diverse and includes prosimians, New World monkeys, Old World monkeys, apes, and humans. More than 250 species and more than 600 subspecies are recognized, and new species are described nearly every year. Recently, Colin Groves has proposed a revised taxonomic system that includes over 300 primate species (Groves, C. 2001. Primate Taxonomy. Washington, DC: Smithsonian Institution Press). The challenge of describing the nutritional needs of primates, which range in size from tiny mouse lemurs and pygmy marmosets to the markedly larger gorillas and orangutans, is daunting, particularly because studies of feeding ecology, gastrointestinal anatomy, and nutrient requirements have been completed for only a few of them. Consequently, data have been sought on one or more model species in eight categories (the suborder Strepsirrhini; the families Hominidae and Pongidae, Hylobatidae, Cercopithecidae, Cebidae, Callitrichidae, and Tarsiidae; and the subfamily Colobinae) in the hope that such data would be representative of the Order. Little information was found on Tarsiidae and Hylobatidae. Over 500,000 primates live in biomedical research laboratories and conservation institutions throughout the world. Records of the regional primate research centers provided by Leo Whitehair of the National Institutes of Health National Center for Research Resources indicate that 16,820 nonhuman primates of 28 species were present in seven U.S. centers at the end of 1998. In 1999, an eighth U.S. center housing 3,638 animals, including about 3,200 baboons, was added. Records of the International Species Information System (at the Minnesota Zoological Garden, Apple Valley, MN; www.worldzoo.org) indicate that over 9,500 nonhuman primates of 145 species were in U.S. and Canadian zoos at the end of 2000. Additional nonhuman primates can be found in U.S. and Canadian government, university, and commercial laboratories. Many primate species serve as surrogates in studies of human physiology and disease, and their nutritional status is known to influence susceptibility and tissue responses to infective agents. The validity of such research is open to question if the experimental subjects have not been appropriately nourished. Likewise, the health and reproduction of primates in zoos can be compromised to an extent that renders the maintenance or multiplication of endangered species impossible. In preparing this report, the Committee was limited in the amount of reliable and specific information available on nutrient requirements, deficiencies, and toxicities in primates. The authors of this publication had as their primary objective the development of guidelines that would ensure that nutrient deficiencies or toxicities and inappropriate dietary husbandry would not limit success in primate research colonies or zoos. We hope that this objective has been fulfilled, in light of the limits of the information available to us, and that researchers will continue to fill the obvious information gaps so that future editions will be more complete. DUANE E. ULLREY, Chair Ad Hoc Committee on Nonhuman Primate Nutrition

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Acknowledgments The Committee wishes to thank the numerous people who provided input by letter or at public forums. In addition, we thank those who took time to meet with the Committee throughout the study process. The financial support provided by the National Center for Research Resources of the National Institutes of Health, the American Zoo and Aquarium Association, The Geraldine R. Dodge Foundation, the Association of Primate Veterinarians, Harlan Teklad, Purina Mills, Inc., and ZuPreem, is gratefully acknowledged. This report has been reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the NRC’s Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the institution in making its published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their review of this report: David J. Baer, US Department of Agriculture Agricultural Research Service, Beltsville, Maryland; Ellen Dierenfeld, The Wildlife Conservation Society, Bronx, New York; Joseph W. Kemnitz, Wisconsin National Primate Research Center, Madison, Wisconsin; Joe Knapka, National Institutes of Health (retired); Terry L. Maple, Zoo Atlanta, Atlanta, Georgia; and Wilson G. Pond, Cornell University, Ithaca, New York. Although the reviewers listed above have provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations nor did they see the final draft of the report before its release. The review of this report was overseen by R. Lee Baldwin, University of California, Davis. Appointed by the National Research Council, he was responsible for making certain that an independent examination of this report was carried out in accordance with institutional procedures and that all review comments were carefully considered. Responsibility for the final content of this report rests entirely with the authoring committee and the institution. Finally, the Committee wishes to thank Charlotte Kirk Baer, program director, Committee on Animal Nutrition, for her encouragement and cheerful guidance of this project to completion. Her exceptional organizational skills contributed in a major way to the success of the Committee. Appreciation also is extended to Stephanie Padgham, project assistant, for her regular communications and helpful provision of supplementary materials.

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Contents     OVERVIEW   1 1   FEEDING ECOLOGY, DIGESTIVE STRATEGIES, AND IMPLICATIONS FOR FEEDING PROGRAMS IN CAPTIVITY,   5     Feeding Ecology,   5     Feeding-Ecology, Methods Involving Visual Observations of Behavior,   5     Observation Options,   5     Sampling Methods,   5     Alternative Feeding-Ecology Methods,   13     Analysis of Stomach Contents,   13     Fecal Analysis,   16     Food Remnants,   16     Reporting Feeding Behavior,   18     Feeding Time,   18     Mass of a Food as Percentage of Total Diet Mass,   18     Feeding-Ecology Tables,   18     Plant-Feeding Strategies,   19     Insect Foraging and Feeding,   19     Additional Considerations,   19     How to Use This Information,   20     Digestive Strategies,   20     Faunivores,   21     Frugivores,   21     Folivores,   24     Implications for Feeding Programs in Captivity,   26     References,   27 2   ENERGY   41     Units of Measurement,   41     Classification,   41     Gross Energy,   41     Digestible Energy,   41     Metabolizable Energy,   42     Physiologic Fuel Values,   42     Requirements,   42     Basal Energy Expenditures or Basal Metabolic Rate,   43

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003     Estimating Basal Metabolic Rate,   43     Effects of Age and Body Composition on Basal Metabolic Rate,   43     Energy Requirements for Maintenance,   43     Energy Requirements for Growth,   48     Energy Requirements for Pregnancy and Lactation,   53     References,   54 3   CARBOHYDRATES AND FIBER   58     Carbohydrate Classification, Characteristics, Digestion, and Metabolism,   58     Monosaccharides,   58     Disaccharides,   59     Oligosaccharides,   59     Polysaccharides,   59     Starch and Starch-Like Polysaccharides,   59     Non-Starch Polysaccharides,   59     Analytical Procedures for Carbohydrate and Fiber,   61     Crude Fiber,   61     Total Dietary Fiber,   61     Neutral-Detergent Fiber and Related Fractions,   62     Carbohydrates in Wild Food Plants,   64     Significance of Fiber,   66     Proposed Fiber Intakes by Nonhuman Primates,   68     Fiber Recommendations for Other Species,   68     Fiber in Wild Food Plants as Guides for Captive-Diet Fiber Concentrations,   68     Fiber Digestion by Nonhuman Primates as a Guide for Captive-Diet Fiber Concentrations,   70     Proposed NDF and ADF Concentrations in Captive Nonhuman Primate Diets,   70     References,   70 4   PROTEIN   75     Protein Sources,   75     Assessment of Protein Requirements,   75     Methods,   75     Digestibility,   76     Requirements,   77     Protein Quality,   77     Proteins Limiting in Sulfur Amino Acids,   77     Proteins Limiting in Lysine,   78     Amino Acid Requirements,   78     Lysine and Methionine,   78     Phenylalanine,   79     Tryptophan,   79     Taurine,   79     Efficiency of Protein Use,   79     Protein Deficiency,   80     Protein for Pregnancy and Lactation,   80     Protein-Calorie Malnutrition in Young Primates,   80     Protein Excess,   83     Non-Amino-Acid Effects of Protein Sources,   83     References,   84

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 5   FATS AND FATTY ACIDS   87     Fat Absorption,   88     Milk Fats,   89     Essential n-3 Fatty Acids,   89     Essential n-6 Fatty Acids,   90     Detrimental Fatty Acids,   91     Cholesterol,   91     Primates as Cardiovascular Disease Models,   92     References,   92 6   MINERALS   94     Macrominerals,   95     Calcium and Phosphorus,   95     Magnesium,   97     Potassium,   98     Sodium,   98     Chloride,   98     Sulfur,   98     Trace Minerals,   98     Iron,   98     Copper,   100     Manganese,   101     Zinc,   102     Iodine,   104     Selenium,   104     Cobalt,   106     Chromium,   106     Fluorine,   107     References,   107 7   VITAMINS   113     Fat-Soluble Vitamins,   113     Vitamin A and Carotenoids,   113     Measures of Biologic Activity,   113     Absorption and Circulation of Carotenoids,   114     Vitamin A and Carotenoids in Feedstuffs,   114     Absorption, Circulation, and Storage of Vitamin A,   114     Vitamin A Deficiency,   115     Vitamin A Requirements,   115     Hypervitaminosis A,   115     Vitamin D,   116     Photobiology, Metabolism, and Function of Vitamin D,   116     Measures of Vitamin D Activity,   117     Vitamin D Deficiency,   117     Discrimination Between Vitamin D2 and Vitamin D3 ,   118     Metabolic Resistance to Vitamin D3 in Callitrichids,   118     Animals Not Exposed to Natural Sunlight or Unable to Make Vitamin D in Their Skin,   120     Vitamin D Requirements,   120     Hypervitaminosis D,   121     Vitamin E,   122     Chemistry and Measures of Activity,   122     Absorption, Metabolism, and Excretion,   123

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003     Biologic Functions,   124     Vitamin E Deficiency,   124     Vitamin E Requirements,   125     Vitamin K,   126     Water-Soluble Vitamins,   128     Thiamin,   128     Riboflavin,   129     Pantothenic Acid,   130     Niacin,   131     Vitamin B6,   132     Biotin,   133     Folacin,   134     Vitamin B12,   135     Vitamin C,   137     Choline,   140     Carnitine,   141     Inositol,   141     References,   142 8   WATER   150     Water Content of the Body,   150     Effects of Activity Restriction,   152     Effects of Cold,   152     Effects of Heat and Water Deprivation,   152     Water Sources,   153     Liquid Water Intake,   153     Preformed-Water Intake,   154     Metabolic Water,   154     Water Loss,   154     Water Quality,   155     Water Requirements,   156     References,   157 9   PATHOPHYSIOLOGIC AND LIFE-STAGE CONSIDERATIONS   159     Body Weight,   159     Nutrition from Birth to Weaning,   159     Growth,   159     Mother-Reared Infants,   161     Artificially Reared Infants,   161     Milk Volume and Composition,   161     Volume,   161     Composition of Mother’s Milk,   164     Nutrient Intakes for Milk Replacers,   164     Formulas Used for Artificially Rearing Infant Nonhuman Primates,   165     Long-Term Consequences of Different Modes of Infant Feeding,   166     Weaning Foods and Strategies,   167     Nutrition and Aging,   167     Dietary Restriction,   167     Bone,   170     Immunology,   170     Wound Healing,   170     Atherosclerosis,   171     Body Composition,   171

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003     Obesity,   172     Regulation of Glucose Metabolism,   174     Diabetes,   174     References,   176 10   DIET FORMULATION, EFFECTS OF PROCESSING, FACTORS AFFECTING INTAKE, AND DIETARY HUSBANDRY   182     Diet Formulation,   182     Natural Dietary Habits,   182     Digestive System Structure and Physiology,   182     Nutrient Requirements,   182     Feedstuffs,   182     Diet Formulation,   182     Effects of Processing,   183     Factors Affecting Intake,   184     Influence of Visual, Olfactory, Taste, and Tactile Clues on Food Acceptance,   185     Regulation of Food Intake,   185     Dietary Husbandry,   186     Primary Food Source,   186     Supplements,   186     Browse,   187     References,   188 11   NUTRIENT REQUIREMENTS   191     References,   194 12   COMPOSITION OF FOODS AND FEED INGREDIENTS   195     References,   195 13   FOOD AS A COMPONENT OF ENVIRONMENTAL ENHANCEMENT   259     Goal of Environmental Enhancement,   259     Role of Food and Foraging,   259     Wild Environment versus Captivity,   260     Species Differences,   261     Manipulation of Foraging Opportunities,   261     Live Prey,   262     Exudates and Gums,   262     Water,   262     Higher-Fiber Foods,   263     Epilogue,   263     References,   263     APPENDIX   266     ABOUT THE AUTHORS   269     INDEX   273

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Tables and Figures TABLES 1-1   Prosimian feeding ecology,   6 1-2   Callithrix feeding ecology,   9 1-3   Cebid feeding ecology,   10 1-4   Colobine feeding ecology,   12 1-5   Non-colobine cercopithecine feeding ecology,   14 1-6   Ape feeding ecology,   17 1-7   Form of foregut in genera of subfamily Colobinae,   24 1-8   Examples of food consumed by primates in zoos and in the wild,   27 2-1   Estimated daily metabolizable energy (ME) requirements (as multiples of BMR) for adult captive animals,   45 2-2   Biologic and metabolic parameters of species fed dry diets,   49 2-3   Biologic and metabolic parameters for the young of various species fed liquid or dry diets,   51 3-1   Common dietary carbohydrates and their digestion,   61 3-2   Fiber concentrations in wild-primate diets (% of dry matter) in studies in which over 70% of items were analyzed,   65 3-3   Fiber concentrations in wild-primate diets (% of dry matter) in studies in which under 70% of items were analyzed,   67 3-4   Fiber levels (% of dietary dry matter) fed to primates in captivity,   69 3-5   Proposed fiber concentrations in total dietary dry matter of extruded diets for primate species grouped by relative ability to utilize plant cell wall,   70 4-1   Estimated protein requirements for primates using high-quality reference proteins,   76 4-2   Potency of common proteins measured by bioassay in primates,   78 5-1   Common names, scientific names, and short-form designations of fatty acids,   88 7-1   Survey of data used to estimate vitamin E requirements,   127

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 7-2   Estimates of thiamin requirement,   130 7-3   Estimates of riboflavin requirement,   131 7-4   Estimates of vitamin B6 requirement,   133 7-5   Estimates of folacin requirement,   136 7-6   Estimates of ascorbic acid requirement,   139 9-1   Body weight of captive adult primates,   160 9-2   Body weight of captive primates at various stages of development,   162 9-3   Primate species identified as potentially at increased risk of obesity in captive environments,   164 9-4   Proximate composition of milk from several primate species,   165 9-5   Composition of nonhuman-primate milk, human milk, and human-infant formula,   166 9-6   Physical characteristics of control (ad libitum-fed) and diet-restricted (30 percent restriction) Macaca mulatta after 4.5 years,   171 9-7   Body fat (%) determined with three methods in Western lowland gorillas,   172 10-1   Plant species used in feeding captive primates,   187 11-1   Estimated nutrient requirements of primate model species fed purified or semipurified diets,   192 11-2   Estimated adequate nutrient concentrations in diets containing conventional feed ingredients intended for post-weaning nonhuman primates, accounting for potential differences in nutrient bioavailabilities and adverse nutrient interactions, but not accounting for potential losses in feed processing and storage,   193 12-1   Composition of important feeds: Energy values, proximate analyses, plant cell wall constituents, data expressed as-fed and dry (100% dry matter),   197 12-2   Composition of important feeds: Minerals, data expressed as-fed and dry (100% dry matter),   213 12-3   Composition of important feeds: Vitamins, data expressed as-fed and dry (100% dry matter),   228 12-4   Composition of important feeds: Amino acids, data expressed as-fed and dry (100% dry matter),   242 12-5   Mineral concentrations in macromineral sources,   256 12-6   Characteristics and energy values of various sources of fats and oils (data on as-fed basis),   258 A-1   Taxonomic relationships, genera, and partial list of species in Order Primates, based on Napier and Napier (1985), Oates et al. (1989), and Nowak (1999),   266 A-2   Weight equivalents,   268 A-3   Weight-unit conversion factors,   268

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 FIGURES 1-1   Gastrointestinal Tract of Tarsier,   22 1-2   Gastrointestinal Tract of Squirrel Monkey,   22 1-3   Gastrointestinal Tract of Night Monkey,   22 1-4   Gastrointestinal Tract of Woolly Monkey,   22 1-5   Gastrointestinal Tract of Vervet Monkey,   23 1-6   Gastrointestinal Tract of Macaque,   23 1-7   Gastrointestinal Tract of Baboon,   23 1-8   Gastrointestinal Tract of Bush Baby,   23 1-9   Gastrointestinal Tract of Northern Douc Langur,   25 1-10   Gastrointestinal Tract of Colobus Monkey,   25 1-11   Gastrointestinal Tract of Chimpanzee,   25 1-12   Gastrointestinal Tract of Orangutan,   25 1-13   Gastrointestinal Tract of Howler Monkey,   26 1-14   Gastrointestinal Tract of Adult Human,   26 3-1   Plant Cell Components in the Analytical Fractions of the Sequential Detergent System of Robertson and Van Soest,   63

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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 Nutrient Requirements of Nonhuman Primates Second Revised Edition, 2003

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