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NUTRIENT REQUIREMENTS OF DOGS AND CATS Ad Hoc Committee on Dog and Cat Nutrition Committee on Animal Nutrition Board on Agriculture and Natural Resources Division on Earth and Life Studies National Research Council

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THE NATIONAL ACADEMIES PRESS 500 Fifth Street, N.W. Washington, DC 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 National Institutes of Health under Agreement No. IR24RR142238-01, the Food and Drug Administration under Agreement No. FD-R001887-01, and the Pet Food Institute. Any opinions, findings, con- clusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the views of the organizations or agencies that provided support for the project. Library of Congress Cataloging-in-Publication Data National Research Council (U.S.). Ad Hoc Committee on Dog and Cat Nutrition. Nutrient requirements of dogs and cats / Ad Hoc Committee on Dog and Cat Nutrition, Committee on Animal Nutrition, Board on Agriculture and Natural Resources, Division on Earth and Life Studies, National Research Council.--[Rev. ed.]. p. cm. Includes bibliographical references and index. ISBN 0-309-08628-0 (hardcover)--ISBN 0-309-50733-2 (pdf) 1. Dogs--Nutrition. 2. Cats--Nutrition. 3. Dogs--Food. 4. Cats--Food. I. Title. SF427.4N38 2006 636.7'0852--dc22 2005037411 Additional copies of this report are available from the National Academies Press, 500 Fifth Street, N.W., Lockbox 285, Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet, http://www.nap.edu. Copyright 2006 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distin- guished 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 char- ter 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. Ralph J. Cicerone 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 autono- mous in its administration and in the selection of its members, sharing with the National Acad- emy 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. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to se- cure 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 educa- tion. 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 gen- eral 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 provid- ing 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. Ralph J. Cicerone and Dr. Wm. Wulf are chairman and vice chairman, respectively, of the National Research Council. www.national-academies.org

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AD HOC COMMITTEE ON DOG AND CAT NUTRITION DONALD C. BEITZ, Chair, Iowa State University, Ames JOHN E. BAUER, Texas A & M University, College Station KEITH C. BEHNKE, Kansas State University, Manhattan DAVID A. DZANIS, Dzanis Consulting & Collaborations, Santa Clarita, California GEORGE C. FAHEY, University of Illinois, Urbana-Champaign RICHARD C. HILL, University of Florida, Gainesville FRANCIS A. KALLFELZ, Cornell University, Ithaca, New York ELLEN KIENZLE, Ludwig Maximilians University, Munich, Germany JAMES G. MORRIS, University of California, Davis QUINTON R. ROGERS, University of California, Davis National Research Council Staff JAMIE S. JONKER, Study Director COMMITTEE ON ANIMAL NUTRITION GARY L. CROMWELL, Chair, University of Kentucky, Lexington C. ROSELINA ANGEL, University of Maryland, College Park JESSE P. GOFF, U.S. Department of Agriculture, Agricultural Research Service, Ames, Iowa RONALD W. HARDY, University of Idaho, Hagerman KRISTEN A. JOHNSON, Washington State University, Pullman BRIAN W. MCBRIDE, University of Guelph, Ontario, Canada KEITH E. RINEHART, Perdue Farms, Inc., Salisbury, Maryland L. LEE SOUTHERN, Louisiana State University, Baton Rouge DONALD R. TOPLIFF, West Texas A&M University, Canyon National Research Council Staff CHARLOTTE KIRK BAER, Director JAMIE S. JONKER, Program Officer STEPHANIE PADGHAM, Project Assistant JOE ESPARZA, Project Assistant MARINA PEUNOVA-CONNOR, Project Assistant v

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BOARD ON AGRICULTURE AND NATURAL RESOURCES MAY BERENBAUM, Chair, University of Illinois, Urbana-Champaign SANDRA BARTHOLMEY, University of Illinois, Chicago ROGER N. BEACHY, Donald Danforth Plant Science Center, St. Louis, Missouri H.H. CHENG, University of Minnesota, St. Paul W.R. GOMES, University of California, Oakland ARTURO GOMEZ-POMPA, University of California, Riverside PERRY R. HAGENSTEIN, Institute for Forest Analysis, Planning, and Policy, Wayland, Massachusetts JEAN HALLORAN, Consumer Policy Institute/Consumers Union, Yonkers, New York HANS R. HERREN, Millennium Institute, Arlington, Virginia DANIEL P. LOUCKS, Cornell University, Ithaca, New York WHITNEY MACMILLAN (Emeritus), Cargill, Incorporated, Minneapolis, Minnesota BRIAN W. MCBRIDE, University of Guelph, Ontario, Canada TERRY MEDLEY, E.I. duPont de Nemours and Company, Wilmington, Delaware OLE NIELSEN (Emeritus), Ontario Veterinary College, Guelph, Canada ROBERT PAARLBERG, Wellesley College, Watertown, Massachusetts ALICE N. PELL, Cornell University, Ithaca, New York BOBBY PHILLS, Florida A&M University, Tallahassee SONYA SALAMON, University of Illinois, Urbana-Champaign PEDRO A. SANCHEZ, The Earth Institute at Columbia University, Palisades, New York B.L. TURNER II, Clark University, Worcester, Massachusetts JAW-KAI WANG, University of Hawaii, Manoa TILAHUN D. YILMA, University of California, Davis National Research Council Staff CHARLOTTE KIRK BAER, Director (until October 2004) ROBIN SCHOEN, Director (from November 2004) KAREN IMHOF, Administrative Assistant AUSTIN LEWIS, Program Officer PEGGY TSAI, Research Associate vi

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Preface In 2000, the National Research Council Committee on of Nutrient Requirements of Dogs (1985) and Nutrient Animal Nutrition convened the ad hoc Committee on Requirements of Cats (1986). As the committee delved into Nutrient Requirements of Dogs and Cats to revise the 1985 this task, it became apparent that we needed to examine ear- and 1986 publications on Nutrient Requirements of Dogs lier research as well to formalize the boundaries of knowl- and Nutrient Requirement of Cats, respectively. The task edge for each topic explored in this report. In some in- presented to the committee was as follows: stances--for example, digestive physiology and physical activity--the chapters are essentially new additions requir- An ad hoc committee of the Committee on Animal Nutrition ing a thorough review of literature through many decades. will revise the 1985 and 1986 publications on Nutrient The committee was also cognizant of the various audi- Requirements of Dogs and Nutrient Requirements of Cats. ences for this report. The report will be used by profession- These two publications, originally published as two separate als in industry and academia for formulating diets and iden- documents, will be revised into a single report. The new re- tifying new topics for research. Government officials may port will provide updated estimates of requirements for all use the report as guidance for regulations for pet food label- nutrients and will contain discussions of nutrient metabo- ing. Students and teachers at universities will use the report lism, toxicity, deficiency, and nutritionally related disease in both dogs and cats. Information on impacts of physiologic as a textbook for dog and cat nutrition. Finally, pet owners status, temperature, breed, age, and environment on nutrient will use the report in evaluating feeding decisions for their requirements will also be included. The revised report will pets. With these varied audiences, the committee chose to address unique biological characteristics affecting nutrient err on the side of caution and include adequate detail from digestion and utilization. General considerations regarding the literature cited to provide a clear road map for how the feed ingredients, diet formulation, and feed processing and recommendations were derived. manufacturing will be presented. Principles of feeding pet The reader should note that this published report reflects and laboratory animals will be addressed. a number of changes made to correct and update an unedited prepublication version of the report. Some values, particu- Throughout the study process, the committee sought larly in Chapter 15, have been revised or deleted based on input from various sources. We held public meetings in con- the availability of new information or to correct errors in cal- junction with professional meetings and invited experts to culation. These changes were examined by four independent speak with us as we worked to complete our task. Over the reviewers and have been approved by the authoring com- course of 3 years, the committee held six meetings and four mittee and the institution. public sessions. We acquired data and information from Our assignment was a challenging one. We are pleased to various public and private organizations. By combining a provide what we believe is a comprehensive document that thorough literature review with a critical analysis of scien- will improve both the understanding of dog and cat nutrition tific data and professional experiences, the committee and its practical application in feeding dogs and cats. developed recommendations that are firmly grounded in science. The report is organized into 15 chapters, each addressing unique aspects of nutrition for dogs and cats or providing DONALD C. BEITZ, Chair summary tables of requirements and feed ingredients. Each Ad Hoc Committee on Nutrient chapter examines literature published since the last editions Requirements of Dogs and Cats vii

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Acknowledgments This new volume represents the integrated efforts of Southern, Louisiana State University; C. Edward Stevens, many individuals. The committee thanks everyone who North Carolina State University; Mark Subramanyam, provided input to this study. We thank all those who pro- Schering-Plough Animal Health Corp.; Duane E. Ullrey, vided opportunities for our public meetings and who par- Michigan State University; and Thomas R. Zeigler, Zeigler ticipated in our public sessions. The financial support pro- Bros., Inc.; and Jrgen Zentec, Veterinary University of vided by the National Institutes of Health, Food and Drug Vienna. Four of the reviewers listed above provided com- Administration, and the Pet Food Institute is gratefully ments on changes made to the prepublication draft. acknowledged. Although the reviewers listed above have provided many During the course of its deliberations, the committee constructive comments and suggestions, they were not asked sought assistance from several people who gave generously to endorse the conclusions or recommendations nor did they of their time to provide advice and information that was con- see the final draft of the report before its release. The review sidered in our deliberations. Special thanks are due to Gail of this report was overseen by Joe Knapka, National Czarnecki-Maulden for extraordinary contribution to the Institutes of Health (retired), appointed by the Division on initiation of this study. Earth and Life Studies. This individual was responsible for This report has been reviewed in draft form by individu- making certain that an independent examination of this re- als chosen for their diverse perspectives and technical ex- port was carried out in accordance with institutional proce- pertise in accordance with procedures approved by the dures and that all review comments were carefully consid- National Research Council's Report Review Committee. ered. Responsibility for the final content of this report rests The purpose of this independent review is to provide candid entirely with the authoring committee and the institution. and critical comments that will assist the institution in mak- Finally, the committee wishes to thank Charlotte Kirk ing its published report as sound as possible and to ensure Baer, program director, and Jamie Jonker, program officer, that the report meets institutional standards for objectivity, Committee on Animal Nutrition, for their encouragement evidence, and responsiveness to the study charge. The re- and cheerful guidance of this project to completion. Their view comments and draft manuscript remain confidential to exceptional organizational skills contributed in a major way protect the integrity of the deliberative process. We wish to to the success of the committee. Appreciation also is ex- thank the following individuals for their review of this re- tended to Stephanie Padgham, Joe Esparza, and Marina port: Leonard S. Bull, North Carolina State University; Peunova-Connor, project assistants for the study, for their Wouter Hendriks, Massey University; Kenneth W. Hinch- regular communications and helpful provision of supple- cliff, Ohio State University; Bruce J. Holub, University of mentary materials. Thanks are also due to Karen Imhof, ad- Guelph; Daniel T. Hopkins, Purina Mills, Inc. (retired); ministrative assistant, who incorporated edits into the final Randall A. Johnson, Animal Nutrition Consulting; Neal version of the report, and to Robin Schoen, who took the Merchen, University of Illinois, Urbana-Champaign; committee through its final deliberations. William Rumpler, U.S. Department of Agriculture; L. Lee ix

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Contents OVERVIEW 1 1 COMPARATIVE DIGESTIVE PHYSIOLOGY OF DOGS AND CATS 5 Introduction, 5 Digestive Compartments and Functions, 6 Mouth, 6 Esophagus, 7 Stomach, 7 Gastric Emptying, 8 Small Intestine, Pancreas, and Liver, 8 Large Intestine, 10 Hormonal Aspects of Digestion, 12 Measurement of, and Factors Affecting, Digestibility, 12 References, 17 2 FEEDING BEHAVIOR OF DOGS AND CATS 22 Food Ingestion by Feral Dogs and Cats, 22 Diurnal Rhythms, and Feeding and Drinking Patterns, 22 Role of Imprinting and Social Factors on Feeding Behavior, 23 Dietary Choice, Palatability, Learned Taste Aversions, and Preferences, 24 Control of Food Intake, 25 Feeding Recommendations, 25 References, 26 3 ENERGY 28 Introduction, 28 Energy Evaluation and Requirements: Two Ways of Looking at the Issue, 28 Energy Evaluation, 28 Methods of Estimating Energy in Food, 28 Prediction of Energy Content, 29 Energy Requirements of Dogs, 33 Definitions, 33 Basis for Establishing Energy Requirements, 33 Basal Metabolic Rate, 34 Diet-Induced Thermogenesis and Resting Fed Metabolic Rate, 35 Adult Maintenance, 35 Gestation, 36 xi

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xii CONTENTS Lactation, 37 Growth, 38 Physical Activity, 39 Energy Requirements of Cats, 39 Basis for Establishing Energy Requirements, 39 Basal Metabolic Rate and Dietary Thermogenesis, 40 Adult Maintenance, 40 Gestation, 42 Lactation, 42 Growth, 45 Energy Deficiency and Excess in Dogs and Cats, 45 Body Condition, 45 References, 45 4 CARBOHYDRATES AND FIBER 49 Definition, Classification, and Measurement, 49 Absorbable Carbohydrates, 49 Digestible Carbohydrates, 49 Fermentable Carbohydrates, 49 Nonfermentable Carbohydrates, 51 Absorbable Carbohydrates (Monosaccharides and Sugar Alcohols), 51 Presence in Foodstuffs, 51 Digestion, Absorption, and Utilization, 51 Factors Affecting Digestibility, Absorption, and Utilization, 51 Nutritive Value, 52 Physicochemical Effects, 52 Physiological Effects, 52 Health-Related Characteristics, 54 Digestible Carbohydrates (Disaccharides, Certain Oligosaccharides, and Nonstructural Polysaccharides), 54 Presence in Foodstuffs, 54 Digestion, Absorption, and Utilization, 54 Factors Affecting Digestibility, Absorption, and Utilization, 55 Nutritive Value, 57 Physicochemical Effects, 57 Physiological Effects, 58 Health-Related Characteristics, 59 Fermentable Carbohydrates (Lactose, Certain Oligosaccharides, Dietary Fibers, and Resistant Starch), 60 Presence in Foodstuffs, 60 Digestion, Absorption, and Utilization, 60 Factors Affecting Digestibility, Absorption, and Utilization, 61 Nutritive Value, 62 Physicochemical Effects, 62 Physiological Effects, 63 Health-Related Characteristics, 67 Poorly Fermentable Carbohydrates (Cellulose and Wheat Bran), 69 Presence in Foodstuffs, 69 Digestion, Absorption, and Utilization, 69 Factors Affecting Digestibility, Absorption, and Utilization, 70 Nutritive Value, 70 Physicochemical Effects, 70 Physiological Effects, 70 Health-Related Characteristics, 71 Carbohydrates in Dog and Cat Diet Formulations, 72 References, 74

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CONTENTS xiii 5 FAT AND FATTY ACIDS 81 Importance of Dietary Fats and Fatty Acids, 81 Types and Sources of Dietary Fats and Fatty Acids, 81 Nomenclature, 81 Sources, 82 Trans Fatty Acids, 83 Medium-Chain Triacylglycerols, 83 Conjugated Linoleic Acid, 83 Other Sources, 84 Analytical Procedures, 84 Acid Hydrolysis, 84 Extraction and Saponification of Fatty Acids, 84 Extraction for Total Lipid Analysis, 84 Fat Energy and Nutrient Density, 85 Digestion and Absorption, 85 Preduodenal Lipase, 85 Pancreatic Lipase, 85 Other Lipases, 86 Summary of Triacylglycerol Hydrolysis, 86 Absorption, 86 Digestibility, 87 Biochemical Basis of Fatty Acid Essentiality, 88 Biosynthesis, 88 Desaturation and Elongation, 88 Fatty Acid Ratios, 88 Essential Fatty Acid Utilization for Structure and Function, 90 Comparative Studies, 90 Membrane Composition and Integrity, 90 Lipid-Protein Interactions, 90 Eicosanoid Production, 90 Growth and Development, 91 Canine Studies, 92 Signs of Deficiency or Excess, 92 Skin and Hair Coat, 93 Inflammatory and Immune Cell Structure and Function, 94 Feline Studies, 95 Signs of Deficiency or Excess, 95 Evidence for Limited 6 Desaturase Activities, 96 Dietary LCPUFA and Feline Life Stage, 96 Immune, Inflammatory, and Platelet Cell Functions, 97 Requirements, Recommendations, and Allowances, 97 Essential Fatty Acids for Dogs and Cats, 98 Dogs, 98 Fat Content of Diets: Preferences, Minima, and Maxima, 98 Growth, 99 Adult Maintenance, 100 Gestation and Lactation, 101 Cats, 102 Fat Content of Diets: Preferences, Minima, and Maxima, 102 Growth, 102 Adult Maintenance, 103 Gestation and Lactation, 104 References, 104

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xiv CONTENTS 6 PROTEIN AND AMINO ACIDS 111 Basic Concepts, 111 Introduction, 111 Structure and Function, 111 Essentiality of Amino Acids, 112 Digestibility and Bioavailability of Protein and Amino Acids, 112 Assessing Protein and Amino Acid Status, 114 Requirements, Allowances, and Tolerances of Protein and Amino Acids, 114 Role of Metabolic Adaptation in Protein and Amino Acid Nutrition, 114 Nitrogen (Crude Protein) Minimum Requirements, Recommended Allowances, and Adequate Intakes, 116 Amino Acid Minimum Requirements, Recommended Allowances, Adequate Intakes, and Safe Upper Limits, 120 Dispensable Amino Acids, 134 Amino Acid Imbalances and Antagonisms, 137 References, 138 7 MINERALS 145 Introduction, 145 Minerals and Acid-Base Balance, 145 Macrominerals, 147 Calcium, 147 Phosphorus, 152 Magnesium, 156 Sodium, 159 Potassium, 162 Chloride, 166 Trace Minerals, 168 Iron, 168 Copper, 171 Zinc, 173 Manganese, 177 Selenium, 179 Iodine, 181 Other Minerals, 184 Arsenic, 184 Boron, 184 Chromium, 184 Molybdenum, 185 Silicon, 185 Nickel, 185 Vanadium, 186 Miscellaneous Minerals, 186 References, 186 8 VITAMINS 193 Introduction, 193 Vitamin A, 194 Absorption, Transport, and Storage, 195 Biological Function, 196 Dogs, 197 Cats, 199 Vitamin D, 200 Absorption, Transport, and Storage, 200 Biological Function, 201

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CONTENTS xv Dogs, 201 Cats, 204 Vitamin E, 205 Absorption, 206 Biological Function, 206 Dogs, 208 Cats, 209 Vitamin K, 210 Absorption, Metabolism, and Excretion, 210 Biological Function, 210 Dogs, 211 Cats, 212 Thiamin (Vitamin B1), 212 Absorption, 213 Biological Function, 213 Dogs, 213 Cats, 215 Riboflavin, 216 Absorption, 216 Biological Function, 216 Dogs, 216 Cats, 217 Vitamin B6, 218 Absorption, Transport, and Storage, 218 Biological Function, 218 Dogs, 219 Cats, 220 Niacin, 220 Absorption, 221 Biological Function, 221 Dogs, 222 Cats, 223 Pantothenic Acid, 223 Absorption, Transport, and Storage, 223 Biological Function, 224 Dogs, 224 Cats, 225 Cobalamin (Vitamin B12), 225 Absorption, 225 Biological Function, 226 Dogs, 226 Cats, 227 Folic Acid, 227 Absorption, Transport, and Storage, 228 Biological Function, 228 Dogs, 228 Cats, 229 Biotin, 230 Absorption, 230 Biological Function, 230 Dogs, 231 Cats, 231 Choline, 231 Absorption, 232 Biological Function, 232

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xvi CONTENTS Dogs, 232 Cats, 233 Ascorbic Acid, 234 Biological Function, 234 Dogs, 235 Cats, 235 Vitamin-Like Substances, 235 Vitamin Losses During Processing and Storage of Dog and Cat Foods, 236 References, 237 9 WATER 246 General Principles, 246 Function, 246 Regulation of Intake by Dogs and Cats, 246 Deficiency in Dogs and Cats, 247 Requirements of Dogs, 247 Requirements for Maintenance, 247 Requirements for Other Life Stages, 248 Requirements for Work and Environmental Stress, 248 Requirements of Cats, 249 Requirements for Maintenance and Other Life Stages, 249 Water and Feline Lower Urinary Tract Health, 249 Recommended Allowances, 249 Toxicity in Dogs and Cats, 250 References, 250 10 SPECIAL CONSIDERATIONS FOR LABORATORY ANIMALS 252 Requirements, 252 Energy, 252 Nutrients Other Than Energy, 252 Food Selection, 252 General, 252 Certified Diets, 254 Purified and Chemically Defined Diets, 254 Foods for Specific Dietary Purposes (e.g., Medical Foods), 254 Feeding Management, 255 Life-Stage Designation of Food, 255 Form of Food, 255 Amount Fed, 255 Frequency of Feeding, 256 Storage of Food, 256 References, 256 11 PHYSICAL ACTIVITY AND ENVIRONMENT 258 Introduction, 258 Calculations and Assumptions, 258 How Much Exercise Do Dogs and Cats Undertake?, 259 Dogs, 259 Cats, 260 Types of Exercise: Sprinting Versus Endurance, 260 Sprint Exercise Over a Distance of Less Than One Kilometer, 260 Endurance Exercise Over Distances Greater Than One Kilometer, 260 Traditional Diets Fed to Racing Dogs, 260 Sled Dogs, 260 Greyhounds, 262

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CONTENTS xvii Diet and Health During Exercise, 262 Effect of Exercise on Intestinal Function, 262 Exercise and Digestibility, 263 When to Feed Exercising Dogs, 263 Exercise and Body Condition, 265 Effect of Confinement and Training, 266 Energy Requirements, 267 Minimal Metabolism, 267 Anxiety, 267 Circadian Rhythms, 268 Cold-Induced Thermogenesis, 268 Nutrients in a Warm Environment, 272 Effect of Reduced Oxygen at High Altitude, 272 Effect of Exercise on Energy Requirements, 273 Effect of Exercise on Energy Required in the Absence of Exercise, 273 Energy Required for Exercise: Dynamic Exercise (Short Runs) Versus Steady-State Exercise (Long Runs), 273 Maximal Oxygen Consumption, 279 Intensity of Exercise, 281 Field Metabolic Rate, 281 Nutrient Requirements, 284 How May Nutrient Requirements Change with Exercise or in the Cold?, 284 Sources of Energy: Fat Versus Carbohydrate, 285 Protein and Exercise, 287 Studies That Have Altered Both Fat and Protein in the Diet of Exercising Dogs, 291 Dietary Carbohydrate and Exercise, 292 Importance of Water and Water Requirements at High Ambient Temperatures, 292 Exercised-Induced Hyperthermia and the Importance of Water During Exercise, 293 Water Requirements at Rest and During Exercise, 294 Sodium, Potassium, and Chloride, 296 Fluids Containing Electrolytes, 298 Calcium, Phosphorus, and Magnesium, 298 Trace Minerals, 299 Vitamins, 300 Other Nutrients, 303 References, 304 12 DIET FORMULATION AND FEED PROCESSING 313 Petfood Processing, 313 General Considerations, 313 Dogs, 313 Cats, 313 Petfood Categories, 313 Dry Petfoods, 313 Semi-moist Petfoods, 314 Soft-Expanded (Soft Dry) Petfoods, 315 Baked Petfoods, 315 Starch-Lipid Complexes, 315 Snacks for Pets, 315 Canned (Wet) Petfoods, 316 Homemade Diets, 317 Diet Formulation, 317 Summary, 317 References, 318

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xviii CONTENTS 13 NUTRIENT COMPOSITION OF INGREDIENTS USED IN DOG AND CAT FOODS 319 References, 343 14 OTHER FOOD CONSTITUENTS 344 Constituents Intended for Nutritive Value, 344 Introduction, 344 Chondroprotective Agents, 344 Antioxidants, 345 Probiotics (Direct-Fed Microbials) and Enzymes, 347 Herbs and Botanicals, 347 Other, 348 Constituents Intended for Technical or Nonnutritive Effects on Food, 348 Introduction, 348 Preservatives, 348 Flavors and Extracts, 350 Colors, 350 Other Additives, 351 References, 351 15 NUTRIENT REQUIREMENTS AND DIETARY NUTRIENT CONCENTRATIONS 354 Use of Nutrient Requirement Tables, 355 Recommendations Relative to ME Compared with Requirements Relative to DM, 355 Requirements Relative to ME in the Food Compared with Requirements Relative to BW, 355 Note on Bioavailability, 356 References, 370 APPENDIXES A ABOUT THE AUTHORS 371 B ACRONYMS AND ABBREVIATIONS 374 INDEX 377

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Tables and Figures TABLES 1-1 Gastrointestinal Hormone Characteristics of Dogs and Cats, 13 3-1 Predictive Equations for Metabolizable Energy in Dog Food, 31 3-2 Predictive Equations for Metabolizable Energy in Cat Food, 31 3-3 Basal Metabolic Rate in Dogs, 34 3-4 Reported Maintenance Energy Requirements of Dogs in Relation to Breed, Age, Housing, and Activity, 37 3-5 Recommendations for Growth of Large- and Giant-Breed Dogs, 39 3-6 Reported Daily Maintenance Energy Requirements of Cats, 41 3-7 Body Condition Scoring System, 43 4-1 Summary of Physicochemical, Physiological, and Health-Related Effects of Absorbable, Digestible, Fermentable, and Nonfermentable Carbohydrates in Dog and Cat Nutrition, 53 4-2 Starch Content of Some Starch-Rich Foodstuffs and By-products Used in Dog and Cat Foods, 54 4-3 Influence of Diet Type on Amylase Activity in Chyme of Adult Dogs, 57 4-4 Average Activity of Disaccharidases in Small Intestinal Mucosa of Cats in Relation to Age, 57 4-5 Molecular Structure and Chemical Linkages of Oligosaccharides with Bifidogenic Effects in Dog and Cat Foods, 60 4-6 Characteristics of Selected Fibers and Their Effects on Intestinal Transit Time and Fecal Characteristics of Dogs and/or Cats, 63 4-7 Characteristics of Selected Fibers and Their Effects on Nutrient Digestibility by Dogs and/or Cats, 65 4-8 Characteristics of Selected Carbohydrates and Their Effects on Gastrointestinal Health Characteristics of Dogs and/or Cats, 67 4-9 Safe Upper Limits of Selected Carbohydrates for Adult Dog and Cat Maintenance Diets, 73 5-1 List of Abbreviations of Selected Fatty Acids and Fatty Acid Terminology, 97 6-1 Plasma Amino Acid Concentrations of Kittens, 115 6-2A Plasma Amino Acid Concentrations of Puppies, 115 6-2B Plasma Amino Acid Concentrations from Normal, Small-, and Large-Breed Adult Dogs Fed a Variety of Commercial Diets Known to Be Adequate for Maintenance, 115 6-3 Endogenous Urinary Nitrogen Excretion of Animals Fed a Protein-Free Diet, 117 7-1 Clinical Signs of Phosphorus Deficiency in Dogs in Relation to Physiological Status and Dietary Phosphorus Concentration, 153 xix

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xx TABLES AND FIGURES 8-1 Relative Vitamin E Activity of Various Tocopherols in Preventing Fetal Resorption and Hemolysis, 207 8-2 Estimated Minimal Requirement of Vitamin E Needed to Compensate for the Elevated Vitamin Demand Caused by Some Common Unsaturated Fatty Acids, 208 8-3 Recovery of Vitamins and Carotenoids Added to Extruded Petfoods and Percentage Loss on Storage, 237 11-1 Greyhound Race Distances and Approximate Fastest Times at Tracks in the United States, 260 11-2 Distances and Reported Times for Some Long-Distance Races, 261 11-3 Analyses of Diets Fed to Working Sled Dogs in the Antarctic, Mid-Twentieth Century, 261 11-4 Effect of Changes in Ambient Temperature on Energy Utilization in Acclimatized and Unacclimatized Cats, 271 11-5 Rate of Energy Utilization in Greyhounds, 275 11-6 Cost of Standing and Running During Steady-State Exercise on a Treadmill for Dogs and Cats, 277 11-7 Efficiency of Gaining and Losing Height When Running, 278 11-8 Maximal Oxygen Consumption in Dogs, 280 11-9 Field Metabolic Rate in Dogs, 282 11-10 Recommendations for Metabolizable Energy Requirements of Exercising Dogs, 284 11-11 Adequate Intake of Amino Acids for Greyhounds, 290 11-12 Daily Water Loss from Hydrated and Dehydrated Dogs and Cats as Affected by Temperature, 293 12-1 Nutrient Content of Dry, Semi-moist, and Canned Dog Foods, 317 13-1 Proximate Analysis of Selected Feed Ingredients, 320 13-2 Carbohydrate and Lignin Concentrations of Some Common Ingredients in Canine and Feline Foods, 324 13-3 Total Fat Concentration and Fatty Acid Composition of Selected Feed Ingredients, 326 13-4 Fatty Acid Composition of Selected Fats and Oils, 328 13-5 Amino Acid Composition of Selected Feed Ingredients, 330 13-6 Mineral Content of Selected Ingredients, 334 13-7 Vitamin Content of Selected Ingredients, 338 13-8 Composition of Selected Inorganic Macro-mineral Sources Used in Petfood, 342 13-9 Composition of Selected Inorganic Micro-mineral Sources Used in Petfood, 343 15-1 Change in Requirements Relative to ME if Requirements Vary Directly with BW, 356 15-2 Daily Metabolizable Energy Requirements for Growth of Puppies After Weaning, 356 15-3 Nutrient Requirements for Growth of Puppies After Weaning, 357 15-4 Daily Metabolizable Energy Requirements for Adult Dogs at Maintenance, 359 15-5 Nutrient Requirements of Adult Dogs for Maintenance, 359 15-6 Daily Metabolizable Energy Requirements for Bitches in Late Gestation, 361 15-7 Daily Metabolizable Energy Requirements for Lactating Bitches Based on Number of Puppies and Weeks of Lactation, 361 15-8 Nutrient Requirements of Bitches for Late Gestation and Peak Lactation, 362 15-9 Daily Metabolizable Energy Requirements of Growth in Kittens After Weaning, 364 15-10 Nutrient Requirements for Growth of Kittens After Weaning, 364 15-11 Daily Metabolizable Energy Requirements for Adult Cats at Maintenance, 366 15-12 Nutrient Requirements of Adult Cats for Maintenance, 366 15-13 Daily Metabolizable Energy Requirements for Lactating Queens, 368 15-14 Nutrient Requirements of Queens in Late Gestation and Peak Lactation, 368

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TABLES AND FIGURES xxi FIGURES 1-1 Dog gastrointestinal tract, 6 1-2 Cat gastrointestinal tract, 6 3-1 Principles of bioavailability of energy in animals, 29 3-2 Comparison of experimentally determined and predicted ME values in prepared cat food, 32 3-3 Comparison of experimentally determined and predicted ME values in prepared cat food, 32 3-4 Schematic comparison of body weight changes of queen and bitch during gestation and lactation, 43 5-1 Chemical structure of the n-3 and n-6 polyunsaturated fatty acid families, 82 5-2 Predominant pathways of essential fatty acid metabolism in mammals, 89 5-3 Generalized pathways of eicosanoid formation from n-3 and n-6 PUFAs, 91 6-1 Effect of dietary crude protein and ratio of essential amino acid nitrogen to total amino acid nitrogen (E:T) on weight gain of kittens, 118 11-1 Effect of environmental temperature (C) on energy requirement, 268 11-2 Changes in oxygen consumption before and after a short bout of submaximal exercise, 274 11-3 Cost of running on a treadmill, 276

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