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Committee on Nutrient Requirements of Fish and Shrimp Board on Agriculture and Natural Resources Division on Earth and Life Studies

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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 a grant from the Agricultural Research Service of the United States Department of Agriculture under Contract No. 59-0790-5-186, the National Oceanic and Atmospheric Administration under Award No. NA08OAR4170833; the United Soybean Board under Project No. 8490; and internal National Research Council funds derived from sales of publications in the Animal Nutrition Series. Any opinions, findings, conclusions, 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 Nutrient requirements of fish and shrimp / Committee on the Nutrient Requirements of Fish and Shrimp, Board on Agriculture and Natural Resources, Division on Earth and Life Studies, National Research Council of the National Academies. p. cm. Includes bibliographical references and index. ISBN-13: 978-0-309-16338-5 (cloth) ISBN-10: 0-309-16338-2 (cloth) ISBN-13: 978-0-309-16339-2 (pdf) ISBN-10: 0-309-16339-0 (pdf) 1. Fishes—Nutrition—Requirements. 2. Shrimps—Nutrition—Requirements. 3. Fishes— Feeding and feeds. 4. Shrimps—Feeding and feeds. I. National Research Council (U.S.). Committee on the Nutrient Requirements of Fish and Shrimp. SH156.N865 2011 595.3’88—dc22 2011008752 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 2011 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 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. 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 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. Charles M. Vest is president of the Na - tional 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 Na - tional 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. Ralph J. Cicerone and Dr. Charles M. Vest are chair and vice chair, respectively, of the National Research Council. www.national-academies.org

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COMMITTEE ON NUTRIENT REQUIREMENTS OF FISH AND SHRIMP RONALD W. HARDY, Chair, University of Idaho, Hagerman DELBERT M. GATLIN, III, Vice-Chair, Texas A&M University, College Station DOMINIQUE P. BUREAU, University of Guelph, Ontario LOUIS R. D’ABRAMO, Mississippi State University, Mississippi State D. ALLEN DAVIS, Auburn University, Auburn, Alabama JOHN E. HALVER, University of Washington, Seattle ÅSHILD KROGDAHL, Norwegian School of Veterinary Science, Oslo, Norway FRANÇOISE MÉDALE, French National Institute for Agricultural Research (INRA), Pee-Sur-Nivelle, France SHI-YEN SHIAU, National Taiwan Ocean University, Keelung DOUGLAS R. TOCHER, University of Stirling, Scotland Staff AUSTIN J. LEWIS, Study Director RUTHIE S. ARIETI, Research Associate ERIN P. MULCAHY, Senior Program Assistant (through August 2010) External Support PAULA T. WHITACRE (Full Circle Communications), Editor v

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BOARD ON AGRICULTURE AND NATURAL RESOURCES NORMAN R. SCOTT, Chair, Cornell University, Ithaca, New York PEGGY F. BARLETT, Emory University, Atlanta, Georgia HAROLD L. BERGMAN, University of Wyoming, Laramie RICHARD A. DIXON, Samuel Roberts Noble Foundation, Ardmore, Oklahoma DANIEL M. DOOLEY, University of California, Oakland JOAN H. EISEMANN, North Carolina State University, Raleigh GARY F. HARTNELL, Monsanto Company, St. Louis, Missouri GENE HUGOSON, Minnesota Department of Agriculture, St. Paul KIRK C. KLASING, University of California, Davis VICTOR L. LECHTENBERG, Purdue University, West Lafayette, Indiana PHILIP E. NELSON, Purdue University, West Lafayette, Indiana ROBERT PAARLBERG, Wellesley College, Watertown, Massachusetts KEITH PITTS, Curragh Oaks Consulting, Fair Oaks, California CHARLES W. RICE, Kansas State University, Manhattan HAL SALWASSER, Oregon State University, Corvallis PEDRO A. SANCHEZ, The Earth Institute, Columbia University, Palisades, New York ROGER A. SEDJO, Resources for the Future, Washington, DC KATHLEEN SEGERSON, University of Connecticut, Storrs MERCEDES VÁZQUEZ-AÑÓN, Novus International, Inc., St. Charles, Missouri Staff ROBIN A. SCHOEN, Director KAREN L. IMHOF, Administrative Assistant AUSTIN J. LEWIS, Senior Program Officer EVONNE P. Y. TANG, Senior Program Officer PEGGY TSAI, Program Officer CAMILLA YANDOC ABLES, Associate Program Officer KARA N. LANEY, Associate Program Officer RUTH S. ARIETI, Research Associate JANET M. MULLIGAN, Research Associate KAMWETI MUTU, Research Associate ERIN P. MULCAHY, Senior Program Assistant (through August 2010) KATHLEEN REIMER, Program Assistant (from August 2010) vi

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Acknowledgments This report has been reviewed in draft form by persons to endorse the conclusions or recommendations, nor did chosen for their diverse perspectives and technical expertise they see the final draft of the report before its release. The in accordance with procedures approved by the National review of this report was overseen by Robert R. Stickney, Research Council’s Report Review Committee. The purpose Texas A&M University. Appointed by the National Research of this independent review is to provide candid and criti- Council, he was responsible for making certain that an cal comments that will assist the institution in making its independent examination of this report was carried out in published report as sound as possible and to ensure that the accordance with institutional procedures and that all review report meets institutional standards of objectivity, evidence, comments were carefully considered. Responsibility for the and responsiveness to the study charge. The review com- final content of this report rests entirely with the author com - ments and draft manuscript remain confidential to protect mittee and the institution. the integrity of the deliberative process. We wish to thank The Committee on Nutrient Requirements of Fish and the following for their review of this report: Shrimp thanks the U.S. Department of Agriculture, the National Oceanic and Atmospheric Administration, and the Geoff L. Allan, Port Stephens Fisheries Centre, New United Soybean Board for contributing funds to support the South Wales, Australia committee’s work. The committee has been fortunate to have Ian P. Forster, Fisheries and Oceans Canada, British Dr. Austin Lewis, Senior Program Officer, Ruthie Arieti, Columbia, Canada Research Associate, and Erin Mulcahy, Senior Program As- Menghe H. Li, Mississippi State University, Mississippi sistant, assigned to the committee. Dr. Lewis has provided State excellent guidance, advice, and encouragement throughout Ingrid Lupatsch, Centre for Sustainable Aquaculture the development of the report, and the committee is grate- Research, Swansea University, UK ful for his sustained support and friendship. Ms. Arieti has Wing Keong Ng, Universiti Sains, Malaysia been extremely effective at keeping the process of writing, Marty Alan Riche, Harry K. Dupree Stuttgart National revising, and editing sections moving along smoothly as well Aquaculture Research Center, Agricultural Research as keeping committee members informed and on track. The Service, U.S. Department of Agriculture, FL committee thanks Robin Schoen, Director of the Board on Michael B. Rust, National Marine Fisheries Service, Na- Agriculture and Natural Resources, for her efforts to get the tional Oceanic and Atmospheric Administration, WA revision underway and for her support and encouragement Wendy M. Sealey, Bozeman Fish Technology Center, during its preparation. The committee members wish to U.S. Fish and Wildlife Service, MT thank their colleagues Arlene Ali, Brett Glencross, Katheline Albert G. J. Tacon, Aquaculture Consultant, Vista, CA Hua, Kyeong-Jun Lee, Yu-Hung Lin, Biswamitra Patro, and Carl D. Webster, Aquaculture Research Center, Ken- Guillaume Salze, whose assistance was essential to complete tucky State University, Frankfort the publication and Victoria Blondin for producing original Robert P. Wilson, Emeritus, Mississippi State University, drawings of fish anatomy. Finally, the committee wishes to Mississippi State thank the National Research Council for giving the members Thomas R. Zeigler, Zeigler Bros, Inc., PA the opportunity to produce a new revised version of the re- port that they hope will guide the aquaculture feed industry, Although the reviewers listed above have provided many scientists, students, and others who share their passion for constructive comments and suggestions, they were not asked fish and shrimp nutrition for many years to come. vii

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Contents SUMMARY 1 1 INTRODUCTION 4 References, 5 2 BASIC CONCEPTS AND METHODOLOGY 6 Determination of Nutrient Requirements, 6 Experimental Design and Conditions, 6 Replicates in the Experimental Design, 9 Measured Responses, 9 Estimating Quantitative Nutrient Requirements, 11 Conclusions, 13 References, 14 3 DIGESTIVE PHYSIOLOGY OF FISH AND SHRIMP 15 Fish, 15 Shrimp, 25 Conclusions, 28 References, 29 4 DIETARY ENERGY UTILIZATION AND METABOLIC INTEGRATION 34 Standard Energy Partitioning Scheme—NRC 1981 Nomenclature, 34 Gross Energy and Intake of Energy, 35 Fecal Energy Losses—Digestible Energy, 36 Nonfecal Losses—Metabolizable Energy, 36 Surface Energy Losses, 37 Heat Losses, 37 Basal/Minimal Metabolism, 38 Effect of Body Weight on Basal Metabolism, 38 Effect of Temperature on Basal Metabolism, 39 Basal Metabolism of Shrimp, 40 Maintenance Energy Requirement, 40 Heat Losses for Voluntary Activity, 42 Heat Increment of Feeding, 42 Estimates of Heat Increment of Feeding, 42 Digestion and Absorption Processes, 43 Formation and Excretion of Metabolic Waste, 44 Transformation of Substrates and Retention in Tissues, 44 ix

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x CONTENTS Practical Net Energy Systems, 44 Recovered Energy, 46 Reproduction and Gonads—Ovum Energy, 47 Calculation of Energy Requirement for Growth, 48 Limitations of Nutritional Energetics Approaches, 50 References, 51 5 PROTEINS AND AMINO ACIDS 57 Proteins and Amino Acids: Biochemistry, Roles, and Overview of Metabolism, 57 Essential Amino Acids—Biochemistry, Roles, and Deficiency Signs, 63 Quantitative Protein and Essential Amino Acid Requirements, 70 Quantifying Essential Amino Acid Requirements, 73 Summary of Published Estimates of Essential Amino Acid Requirements of Fish and Shrimp, 75 Essential Amino Acid Requirements in the Context of Feed Formulation, 75 References, 92 6 LIPIDS 102 Fatty Acid Structure and Nomenclature, 102 Lipid Class Structures, 102 General Lipid Metabolism, 105 Dietary Lipid Level, 106 Specific Requirements, 107 Other Issues in Lipid Nutrition, 123 References, 125 7 CARBOHYDRATES AND FIBER 135 Types of Carbohydrates, 135 Nonstarch Polysaccharides in Fish and Shrimp Diets: Physiological Consequences, 144 Digestibility of Starch, 146 Metabolic Fate of Glucose, 149 Nutritional Role of Digestible Carbohydrates in Fish and Shrimp, 156 References, 157 8 MINERALS 163 Calcium and Phosphorus, 168 Magnesium, 170 Sodium, Potassium, and Chloride, 171 Chromium, 172 Copper, 172 Iodine, 173 Iron, 173 Manganese, 175 Selenium, 175 Zinc, 176 Other Minerals, 176 Sources and Forms, 177 Interactions with Other Dietary Components, 179 References, 179 9 VITAMINS 186 Fat-Soluble Vitamins, 186 Water-Soluble Vitamins, 201

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xi CONTENTS Other Vitamin-Like Compound, 209 References, 210 10 FEED ADDITIVES 221 Antimicrobial Agents, 221 Antioxidants, 221 Binding Agents, 222 Color/Pigmentation Agents, 222 Enzymes, 224 Organic Acids, 225 Feeding Stimulants/Palatability Enhancers, 225 Immunostimulants, 226 Probiotics and Prebiotics, 227 Hormones, 228 References, 228 11 ANTINUTRITIONAL FACTORS AND ADVENTITIOUS TOXINS IN FEEDS 232 Antinutrients in Plant Feedstuffs, 233 Combined Effects of Plant Antinutrients, 242 Antinutrients in Animal Feedstuffs, 243 Adventitious Toxins, 244 Unknown Compounds, 246 Conclusions, 246 References, 246 12 DIGESTIBILITY AND AVAILABILITY 253 Methods Used in Digestibility Determination, 253 Digestibility of Feed Ingredients, 255 References, 268 13 NUTRIENT DELIVERY AND FEEDING PRACTICES 272 Feeding Early Life Stages, 272 Production Diets and Feed Management, 273 Feed Utilization and Fish Growth, 274 Pollution Loading and Waste Management, 281 Conclusions, 282 References, 282 14 LARVAL NUTRITION 286 Digestive Enzymes, 286 Relationship of Larval Stage, Duration of Gut Retention, and Level of Enzyme Activity, 287 Nutritional Enrichment of Live Food, 288 Formulated Diets, 288 Nutrient Requirements, 291 Conclusions, 295 References, 295 15 INGREDIENTS, FORMULATION, AND PROCESSING 299 Feed Ingredients, 299 Feed Formulation, 300 Feed Manufacturing, 301 Feed Quality Assessment, 302 Environmental and Sustainability Concerns, 303 References, 303

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xii CONTENTS 16 REPLACEMENT OF MARINE RESOURCES: EFFECTS OF ALTERNATIVE INGREDIENTS AND STRATEGIES ON NUTRITIONAL QUALITY 304 Limitations to Supply and Use of Marine Resources, Fish Meal, and Fish Oil, 304 Substitution of Fish Meal, 305 Substitution of Fish Oil, 308 Conclusions, 318 References, 318 17 CRITICAL RESEARCH NEEDS 323 Requirements, Delivery, and Interaction of Nutrients, 323 Fish Meal and Fish Oil Alternatives, 324 Diet Formulations and Processing, 324 Nutrigenomic Effects and Metabolism, 325 Conclusions, 325 18 NUTRIENT REQUIREMENTS TABLES 326 19 FEED COMPOSITION TABLES 334 20 COMMON AND SCIENTIFIC NAMES OF SPECIES DISCUSSED IN THIS REPORT 348 APPENDIXES A COMMITTEE STATEMENT OF TASK 351 B ABBREVIATONS AND ACRONYMS 352 C COMMITTEE MEMBER BIOGRAPHIES 358 D RECENT PUBLICATIONS OF THE BOARD ON AGRICULTURE AND NATURAL RESOURCES 361 Policy and Resources, 361 Animal Nutrition Program—Nutrient Requirements of Domestic Animals Series and Related Titles, 362 INDEX 363

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Tables and Figures TABLES 3-1 Digestive Enzymes of the Digestive Tract, 21 4-1 Terminology of Types of Dietary Energy and Energy Budget Components, 35 4-2 Estimate of Maintenance Energy Requirement of Different Fish and Shrimp Species Obtained Through Feeding Trials, 41 4-3 Estimates of Maintenance, Cost of Protein and Lipid Deposition, 45 4-4 Energy and Oxygen Requirements and Expected Feed Efficiency of Rainbow Trout (Oncorhynchus mykiss), 49 4-5 Energy and Oxygen Requirements and Expected Feed Efficiency of European Sea Bass (Dicentrarchus labrax), 49 4-6 Energy and Oxygen Requirements and Expected Feed Efficiency of Asian Sea Bass (Lates calcarifer), 50 5-1 Amino Acid Composition of Different Body Proteins of Animals, 58 5-2 Amino Acid Composition (g/16 g N) of Various Fish and Shrimp Species, 59 5-3 Essential and Nonessential Amino Acids, 59 5-4 Recommended Dietary Protein Levels for Various Fish Species of Commercial Importance (As-Fed Basis), 70 5-5 Recommended Dietary Protein Levels of Different Shrimp Species, 71 5-6 Arginine, 76 5-7 Histidine, 78 5-8 Isoleucine, 78 5-9 Leucine, 79 5-10 Lysine, 80 5-11 Methionine, 82 5-12 Phenylalanine, 84 5-13 Threonine, 85 5-14 Tryptophan, 86 5-15 Valine, 86 5-16 Summary of Studies on Essential Amino Acid Requirements of Shrimp, 87 5-17 Arginine Requirement of Rainbow Trout (Oncorhynchus mykiss) According to Different Modes of Expression, 90 5-18 Dietary Arginine Level Expected to Meet Requirement of Rainbow Trout (Oncorhynchus mykiss), 90 5-19 Ideal Amino Acid Profile for Teleost Fish and Penaeid Shrimp Derived from a Synthetic Review of the Literature, 91 xiii

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xiv TABLES AND FIGURES 5-20 Digestible Essential Amino Acid Requirements (% Diet Dry Matter) Estimated Using a Factorial Model for Rainbow Trout (Oncorhynchus mykiss), 91 5-21 Digestible Essential Amino Acid Requirements (% Diet Dry Matter) Estimated with a Factorial Model for Atlantic Salmon (Salmo salar), 91 6-1 Reported Quantitative Essential Fatty Acid Requirements of Juvenile and Subadult Freshwater and Diadromous Species of Finfish, 109 6-2 Reported Quantitative Essential Fatty Acid Requirements of Larvae and Early Juveniles of Finfish, 110 6-3 Reported Quantitative Essential Fatty Acid Requirements of Juvenile and Subadult Marine Species of Finfish, 111 6-4 Reported Quantitative Essential Fatty Acid Requirements of Shrimp, 113 6-5 Reported Phospholipid Requirements in Juvenile and Larval Shrimp Species, 118 6-6 Reported Quantitative and Qualitative Phospholipid Requirements of Finfish, 119 6-7 Reported Cholesterol/Sterol Requirements of Shrimp and Other Crustaceans, 122 7-1 Carbohydrate Categories, 136 7-2 Carbohydrates in Selected Ingredients Used for Fish and Shrimp Diets, 138 7-3 Starch Content and Characteristics of Starches of Some Selected Ingredients for Fish and Shrimp Diets, 141 7-4 Apparent Digestibility Coefficient of Starch According to the Sources and the Dietary Levels in Different Fish and Shrimp Species, 147 7-5 Effect of Oral, Intraperitoneal, or Intravenous Administration of Different Carbohydrates Sources on Blood Glucose Levels and Return to the Basal Level, 150 7-6 Changes in Liver Glycogen Content with Fasting and Feeding Status, 154 7-7 Effect of Carbohydrate Sources and Levels on Glycogen Content in Different Fish and Shrimp Species, 155 8-1 Minerals and Some of Their Prominent Functions and Deficiency Signs Observed in Fish and Shrimp, 163 8-2 Macromineral Requirements of Fish, 164 8-3 Micromineral Requirements of Fish, 166 8-4 Mineral Requirements of Crustaceans, 167 9-1 Historical Vitamin Diagnostic Signs Reported in Fish and Shrimp, 187 9-2 Historical Vitamin Requirements Estimates for Growing Fish Determined with Chemically Defined Diets in a Controlled Environment, 190 9-3 Historical Vitamin Requirements Estimates for Growing Shrimp Determined with Chemically Defined Diets in a Controlled Environment, 194 9-4 Historical Vitamin C Requirements Estimates for Growing Fish and Shrimp with Chemically Defined Diets in a Controlled Environment, 195 10-1 Xanthophyll Content of Plant Materials and Astaxanthin Content of Animal Products Used in Aquatic Feeds, 223 11-1 Important Antinutrients Present in Some Commonly Used Potential Fish Feed Ingredients, 232 11-2 Adventitious Toxins and Other Undesirable Substances That May Contaminate Fish Feed, 233 12-1 Apparent Digestibility of Protein in Selected Feed Ingredients for Several Fish Species and Shrimp, 258

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xv TABLES AND FIGURES 12-2 Amino Acid Availability and Protein Digestibility Values of Selected Feed Ingredients for Atlantic Salmon, Rainbow Trout, Striped Bass, Channel Catfish, Nile Tilapia, Gilthead Sea Bream, Siberian Sturgeon, Largemouth Bass, Pacu, Rockfish, Yellowtail, Silver Perch, and Pacific White Shrimp, 260 12-3 Apparent Digestibility of Energy in Selected Diet Ingredients of Several Fish Species and Pacific White Shrimp (Litopenaeus vannamei), 264 12-4 Apparent Digestibility of Lipid and Carbohydrate in Selected Feed Ingredients for Fish Species and Pacific White Shrimp (Litopenaeus vannamei), 266 12-5 Phosphorus Availability of Selected Feed Ingredients for Several Fish Species and Pacific White Shrimp (Litopenaeus vannamei), 267 13-1 General Daily Feeding Rates and Frequency Guide for the Production of Channel Catfish, Common Carp, and Nile Tilapia, 279 13-2 Example of Daily Digestible Energy (DE) and Feed Requirement of Rainbow Trout (Oncorhynchus mykiss), 279 13-3 Example of Feed Consumption Rates for Channel Catfish (Ictalurus punctatus), 280 16-1 Fatty Acid Compositions (Percentage of Total Fatty Acids) of Major Vegetable Oils and Animal Fats, 310 16-2 World Oil and Fat Production 2008, 311 16-3 Effect of Complete or Partial Replacement of Dietary Fish Oils by Vegetable Oils on Fatty Acid Compositions (Percentage of Weight) of Total Lipid of Flesh of Salmonids and Marine Fish, 313 18-1 Nutrient Requirements of Freshwater Fish (dry-matter basis), 327 18-2 Nutrient Requirements of Marine Fish (dry-matter basis), 329 18-3 Nutrient Requirements of Shrimp (dry-matter basis), 331 18-4 Partial Summary of Deficiency Signs and Pathologies Associated with Deficiencies of Essential Nutrients (For a More Complete Description, Consult the Chapters on Specific Nutrients—Chapters 5–9), 333 19-1 Typical Dry Matter and Proximate Composition Values for Natural and Chemically Defined Ingredients Commonly Used in Aquatic Animal Feeds (as-fed basis), 335 19-2 Amino Acid Composition of Ingredients (as-fed basis), 337 19-3 Mineral Composition of Ingredients Commonly Used in Aquatic Animal Feeds (as-fed basis), 340 19-4 Vitamin Composition of Ingredients Commonly Used in Aquatic Animal Feeds (as-fed basis), 342 19-5 Fatty Acid (Percentage of Total Fatty Acids) and Cholesterol Composition of Common Animal Fats, Fish Oils, and Vegetable Oils (as-fed basis), 345 19-6 Chemical Composition of Some Purified Feed Ingredients Commonly Used for Aquatic Animal Research (as-fed basis), 347 20-1 Common and Scientific Names of Species Discussed in This Report, 348 FIGURES 2-1 Models for the interpretation of dose-response experiments, 12 3-1 Comparative digestive anatomy of fish, 16 3-2 Organization of internal organs in a generalized fish, 17 3-3 Drawing of stomach and pyloric ceca in Atlantic cod (Gadus morhua), 18 3-4 Anatomy of the digestive tract of shrimp, 26

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xvi TABLES AND FIGURES 4-1 Schematic representation of the energy flow through an animal, 35 4-2 Fasting heat losses of rainbow trout, Oncorhynchus mykiss, 39 4-3 Illustration of the concept of maintenance and fasting heat losses, 40 4-4 Recovered energy and metabolizable energy in rainbow trout, Oncorhynchus mykiss, 41 4-5 Recovered energy and metabolizable energy in Atlantic salmon, Salmo salar, 43 5-1 Relationship between protein mass and live weight of rainbow trout (Oncorhynchus mykiss), 58 5-2 Relationship between water mass and protein mass of rainbow trout (Oncorhynchus mykiss), 58 5-3 Effect of lysine and DE content of the diet on efficiency of lysine utilization of rainbow trout (Oncorhynchus mykiss), 63 5-4 Protein intake per kilogram of live weight gain in different fish and shrimp species, chicken, and swine, 71 5-5 Meeting essential amino acid requirements of rainbow trout (Oncorhynchus mykiss) using three different approaches, 89 6-1 Palmitic (16:0) and oleic (18:1n-9) acids showing the n carbon numbering system, 103 Arachidonic (20:4n-6) and docosahexaenoic (22:6n-3) acids showing the n and Δ 6-2 carbon numbering systems, 103 6-3 The structures of cholesterol and triacylglycerol, 104 6-4 The structures of the main phospholipid classes, 104 6-5 Pathways of biosynthesis of C20 and C22 long-chain polyunsaturated fatty acids, 108 7-1 Structure of the main pentoses and hexoses, 136 The α- and β-anomers of glucose, 137 7-2 7-3 Structure of amylose, 141 7-4 Structure of amylopectin, 142 7-5 Categories of dietary carbohydrates based on current analytical methods, 143 7-6 Structure of cellulose, 143 7-7 Structure of a chitosan unit that composes chitin, 145 7-8 Scheme of glycolysis and gluconeogenesis, 153 13-1 Proximate components of rainbow trout, Oncorhynchus mykiss, 274