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Third Edition Fish Nutrition This Page Intentionally Left Blank Third Edition Fish Nutrition Edited by John E Halver School of Aquatic and Fishery Sciences University of Washington Seattle, Washington and Ronald W Hardy Hagerman Fish Culture Experiment Station University of Idaho Hagerman, Idaho Amsterdam Boston London New York Oxford Paris San Diego San Francisco Singapore Sydney Tokyo This book is printed on acid-free paper ∞ Copyright C 2002, 1989, 1972, Elsevier Science (USA) 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 Requests for permission to make copies of any part of the work should be mailed to: Permissions Department, Harcourt, Inc., 6277 Sea Harbor Drive, Orlando, Florida 32887-6777 COVER IMAGES: Sea Bram and Catfish courtesy of New York SAREP Rainbow trout from Behnke, R J 1992 Native Trout of Western North America, American Fisheries Society Monograph 6, Bethesda, Maryland, USA ACADEMIC PRESS An Imprint of Elsevier Science 525 B Street, Suite 1900, San Diego, California 92101-4495, USA http://www.academicpress.com ACADEMIC PRESS Harcourt Place, 32 Jamestown Road, London, NW1 7BY, UK http://www.academicpress.com Library of Congress Cataloging Number: International Standard Book Number: 0-12-319652-3 PRINTED IN THE UNITED STATES OF AMERICA 02 03 04 05 SB Contents List of Contributors xi Preface Bioenergetics Dominique P Bureau, Sadasivam J Kaushik, and C Young Cho 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 1.10 1.11 1.12 1.13 1.14 1.15 1.16 xiii Introduction History of Nutritional Energetics Energy Exchange in Biological Systems Energy Utilization and Requirements Digestible Energy of Feedstuffs Effect of Biological and Environmental Factors Urinary and Branchial Energy and Metabolizable Energy Factors Affecting Metabolic Waste Output Heat Production Minimal Metabolism Heat Increment of Feeding Digestion and Absorption Processes (HdE) Recovered Energy and Growth Reproduction Integrating and Using Information from Bioenergetics Limitations and Perspectives of Bioenergetics References 14 16 18 21 24 29 35 37 43 47 48 53 54 The Vitamins John E Halver 2.1 2.2 2.3 2.4 2.5 Historical Introduction The Water-Soluble Vitamins The Fat-Soluble Vitamins Other Factors Anemias and Hemapoiesis References 62 66 113 128 130 132 v vi Contents Amino Acids and Proteins Robert P Wilson 3.1 3.2 3.3 3.4 3.5 4.6 4.7 4.8 4.9 Introduction Structures and Biosynthesis Functions Fatty Acids and Dietary Energy Optimal Levels and Ratios of Dietary n-3 and n-6 Polyunsaturated Fatty Acids Dietary Phosphoglycerides: Inositol and Choline Fatty Acid Peroxidation Sources of Lipids for Farmed Fish Feeds Prospects References 182 184 194 201 206 227 232 239 244 246 The Minerals Santosh P Lall 5.1 Introduction 5.2 Essential Minerals for Finfish 5.3 Concluding Remarks References 144 145 151 152 170 175 The Lipids John R Sargent, Douglas R Tocher, and J Gordon Bell 4.1 4.2 4.3 4.4 4.5 Introduction Protein Requirements Qualitative Amino Acid Requirements Quantitative Amino Acid Requirements Other Methods of Estimating Amino Acid Needs References 260 271 300 301 Intermediary Metabolism Konrad Dabrowski and Helga Guderley 6.1 6.2 6.3 6.4 Introduction: Metabolic Circuitry and Control Mechanisms Carbohydrate Metabolism Protein and Amino Acid Metabolism Conclusions References 310 313 333 358 360 Contents Nutritional Physiology Michael B Rust 7.1 7.2 7.3 7.4 7.5 7.6 7.7 7.8 7.9 7.10 7.11 Introduction Gross Juvenile and Adult Anatomy Sensory Organs Food Capture Structures and Organs Digestive Organs Liver Anatomy and Diet Digestive Processes Postabsorptive Transport and Processing Control and Regulation of Digestion Nutritional Physiology in Larval Fish References 368 369 378 389 393 413 415 417 427 428 432 446 Nutritional Pathology Ronald J Roberts 8.1 8.2 8.3 8.4 8.5 8.6 8.7 8.8 8.9 8.10 8.11 8.12 8.13 8.14 8.15 8.16 8.17 8.18 vii Introduction Principles of Nutritional Pathology The Deficiency and Imbalance Diseases Micronutrients Mineral Deficiencies and Imbalances Dietary Mineral Toxicity Mycotoxins Toxic Algae Cottonseeds Senecio Alkaloids Leucaena Toxins Anthropogenic Chemicals Binders Photosensitizers Sekoke Disease Spleen- and Liver-Induced Cataracts Single-Cell Protein Lesions Antibiotic and Chemotherapeutic Toxicity References 454 455 459 464 480 484 489 492 492 492 494 494 494 495 497 498 500 500 500 Diet Formulation and Manufacture Ronald W Hardy and Frederick T Barrows 9.1 Introduction 9.2 Aims and Strategy of Fish Feed Production 506 514 viii Contents 9.3 9.4 9.5 9.6 9.7 10 Feed Ingredients Diet Formulation Diet Manufacture and Storage Ingredient and Diet Evaluation Glossary References Adventitious Toxins Jerry D Hendricks 10.1 Introduction 10.2 Naturally Occurring Toxins in Formulated Fish Rations 10.3 Nonnatural Components and Additives in Formulated Rations 10.4 Summary References 11 630 641 641 Introduction Formulation of Special Feeds Feed Manufacturing Summary References 652 652 661 667 668 Nutrition and Fish Health Delbert M Gatlin III 12.1 12.2 12.3 12.4 12.5 13 602 603 Special Feeds George M Pigott and Barbee W Tucker 11.1 11.2 11.3 11.4 12 515 538 558 578 594 596 Introduction Factors Affecting Fish Health Dietary Components Influencing Fish Health Feeding Practices Affecting Fish Health Concluding Remarks and Research Needs References 672 673 675 694 698 699 Diet and Fish Husbandry Richard T Lovell 13.1 13.2 13.3 13.4 13.5 Introduction Channel Catfish Salmonids Tilapias Penaeid Shrimp References 704 708 720 732 741 753 Contents 14 ix Nutrient Flow and Retention John E Halver and Ronald W Hardy 14.1 14.2 14.3 14.4 14.5 14.6 14.7 14.8 14.9 14.10 14.11 14.12 14.13 14.14 14.15 14.16 Introduction Carbohydrate Metabolism Glycolysis Carbohydrate Synthesis Pentose Phosphate Pathway Glycogenolysis Diet and Carbohydrate Metabolism Lipid Metabolism Odd-Chain-Length Fatty Acid Oxidation Electron Transfer Cascade Amino Acid Metabolism Effect of Diet on Intermediary Metabolism Measuring Protein Accretion and Degradation Intake and Metabolism Sexual Maturity and Metabolism Prospects for Improvement of Protein Retention Efficiency References 756 757 757 759 759 759 760 760 762 763 763 765 766 767 767 768 769 Appendix 771 Index 807 810 Index Catecholamines, glycogen metabolism regulation, 321 Catfish, see Channel catfish CCK, see Cholecystokinin Cellulase, characteristics in digestion, 400 Channel catfish advantages in culture, 708 aeration of ponds, 711, 715 compensatory growth, 719 farming history and features, 708–709, 711 feed consumption and conversion ratios, 713, 719 feeding practices diseased fish, 716 fingerlings, 712 fry, 711 production fish, 712–715 timing during day, 714–715 winter feeding, 715–716 flavor and color optimization of product, 718–719 market, 708 nutrient requirements, see also specific nutrients energy, 716–717 fatty acids, 717 minerals, 718 protein and amino acids, 717–718 vitamins, 718 Chemical coupling, nutritional energetics, Chemical score, protein quality assessment in feeds, 587–588 Chemoreception, see Gustation; Olfaction Chemotherapeutics, toxicity, 500 Chitin, immunostimulation, 692 Chitonase characteristics in digestion, 400 features, 423, 425 Chloride deficiency, 284–285 functions, 284 metabolism, 284 requirement, 285 sources, 285–286 Cholecystokinin (CCK), characteristics in digestion, 430 Cholelithiasis, features, 488 Choline clinical assessment of status, 112 deficiency syndrome, 111, 475–476 discovery, 110 functions, 111 larva requirements, 227–231 requirements, 111 sources and preparations, 112 structure and physical properties, 110–111 Chromium, functions and requirements, 299–300 Chymotrypsin, characteristics in digestion, 398 Citric acid, phosphorous digestibility effects in feeds, 546 Citrinin, manifestations in fish, 627–628 Citrovorum factor, features, 129 C max , see Maximum feed consumption CMS, see Cardiomyopathy syndrome Cobalt, functions and requirements, 299 Collagenase, characteristics in digestion, 399 Conditioning, feed manufacture, 562–563 Conversion feed, characteristics, 542–543 Copper deficiency, 289–290, 482 functions, 289 metabolism, 289 requirement, 290 sources, 290 toxicity, 290, 482 Cottonseed protein supplements, 520 toxins, 492 Crawford, Adair, nutritional energetics contributions, 2–3 Cyanoxcobalamin, deficiency, 476 Cyclopropene fatty acids manifestations in fish, 617–618 sources, 616–617 structures, 616 Cysteine, replacement values for methionine, 164 D Daily instantaneous growth rate (G w ), feed evaluation, 588–589 DDT history of use and persistence, 631 manifestations in fish, 631–632 oral toxicity, 630–631 DE, see Digestible energy DHA, see Docosahexaenoic acid Dicoumarol, vitamin K antagonism, 127–128 Dieldrin history of use and persistence, 633–634 manifestations in fish, 634–635 structure, 633 Diet formulation and manufacture adventitious toxins, see specific toxins calculations in formulation best-buy ingredients, 549–551 Index least-cost feed analysis linear programming, 553–558 simultaneous equations, 551–553 concerns and influences, 507 considerations bioavailability, 539 cost, 538–540 ingredient limits, 548 nutritional requirements, 538 pellet stability, 549 type of diet, 548–549 economic evaluation, 594 evaluation, see Ash; Biological evaluation of feeds; Fiber; Lipid; Protein; Proximate analysis glossary of terms, 584–585 historical perspective composition of early diets, 507–508 dry pelleted feed, 510–513 larval feeds, 514 semimoist pelleted feeds, 509–510 semipurified research feeds, 514 wet–dry feed mixtures, 508–509 wet feed, 508 ingredients fats and oils, 523–525 mineral premixes, 526–527 nomenclature, 515–517 nonnutritive additives antimicrobial agents, 533–535 antioxidants, 535–536 carotenoid supplements, 528–529 enzyme supplements, 531 fiber, 536–537 flavorings and palatability enhancers, 537 hormones, 531–532 overview, 526–527 pellet binders, 527–528 probiotics, 530–531 therapeutics and nonspecific immune stimulants, 529–530 water, 537 protein supplements, 517–522 sources, 515 starch from grains, 522–523 vitamin premixes, 525–526 interdependence of formulation and manufacture, 506–507 larval feed manufacture complex feeds, 577–578 microbound feeds crumbled cake feeds, 574–575 crumbled feeds, 573–574 flake feeds, 574 811 microencapsulated feeds, 576–577 on-size feeds microextrusion marumerization, 575–576 particle-assisted rotational agglomeration, 576 spray beadlets, 576 particle size, 573 manufacture processes conditioning, 562–563 expansion, 563 grinding, 558–560 mixing, 560–562 overview, 558 pelleting cold pellet extrusion, 567 compressed pelleting, 563–565 cooling and drying, 569 crumbling and screening, 569–570 extruded dry pelleting, 565–566 objectives, 563 shipping and storage, 571–572 top-dressing with lipids, 571 universal pellet cooker, 566–567 semipurified diets, 572–573 phase feeding and feed characteristics broodstock feeds, 543–544 conversion feeds, 542–543 fingerling feeds, 543 first feed, 540–541 fry feeds, 541–542 goals, 540 grower feeds, 543 low-pollution feeds, 544–547 product quality feeds, 547–548 recirculation system feeds, 547 seawater transfer feeds, 547 transition feeds, 542–543 production aims and strategy, 514–515 ratios of dietary n-3 and n-6 polyunsaturated fatty acids, see Fatty acid special feeds, see Special feeds Digestible energy (DE), see also Apparent digestible energy evaluation in feeds, 592 factors affecting feeding level and frequency, 16–17 water temperature, 17–18 measurement, 10–11, 13 nutritional energetics, 8–9 requirement determination, 51, 53 values for nutrients, 760, 780–783 Digestible protein (DP), metabolic waste output effects, 22 812 Index Digestive processes chemical processes, 418 enzymatic processes carbohydrases, 422–423, 426 classification, 419 lipases, 421–422 modes of digestion, 418–419 proteases and peptidases, 419–421 table of enzymes, 398–400 overview, 417 physical processes, 418 Digestive tract, see also specific organs digestive enzyme characteristics, 398–400 epithelium, 378 gross anatomy, juveniles and adults carnivores, 369–370 configuration comparisons between fish classes, 369–370 histologic staining, 378 layers of tissue, 393 Dinoflagellate toxins feed contamination, 621 sources, 620–621 Dinogunellin, fish meal contamination, 620 Dioxin, feed contamination, 524 Docosahexaenoic acid (DHA) energy utilization, 194–196 fish requirements, 658 human nutrition value, 200–201 membrane structure and function, 196–198 DP, see Digestible protein E Egg energy content, 47 lipid composition freshwater fish, 222–223 marine fish, 210–211, 229–230 Eicosapentaenoic acid (EPA) eicosanoid production interference, 199–200 energy utilization, 194–196 fish requirements, 658 human nutrition value, 200–201 membrane structure and function, 196–198 Elastase, characteristics in digestion, 398 Endergonic reactions, nutritional energetics, 5–6 Energy partitioning scheme and nomenclature, National Research Council, 2, Energy requirements bioenergetics models, 49–50 fish growth prediction, 50–51 rainbow trout, 51–52 EPA, see Eicosapentaenoic acid Erucic acid manifestations in fish, 612 reduction in feeds, 611–612 sources, 611–612 Erythromycin supplementation, 530 toxicity, 500 Esophagus, structure and function, 392–393 Exergonic reactions, nutritional energetics, 5–6 Expansion, feed manufacture, 563 Eye, teleost anatomy, 379–380 cones, 381, 445 larval development and feeding implications, 445–446 light path, 380–381 polarized light detection, 381–382 rods, 381 ultraviolet vision, 381–382, 445 F Fasting heat production (HEf) basal metabolism, 29, 31 factors affecting body weight, 31–32 temperature, 32–33 measurement, 29–31 species differences, 30–31 Fat, see Fatty acid; Lipid Fatty acid, see also Lipid adventitious toxins cyclopropene fatty acids, 616–618 erucic acid, 611–612 biosynthesis, 189–193 deficiency features, 461–462 egg composition freshwater fish, 222–223 marine fish, 210–211, 229–230 essential fatty acids, 204, 206–207 functions energy production, 194–196 membrane structure and function, 196–201 immune system effects n-3 fatty acids, 683–684 oxidized fatty acid, 682–683 membrane specificity, 183 metabolism, 460, 760–762 optimal levels and ratios of dietary n-3 and n-6 polyunsaturated fatty acids Index essential fatty acid requirements, 206–207 freshwater fish broodstock requirements, 226–227 embryogenesis and early larval requirements, 222–224 juvenile and adult requirements, 225–226 larva and fry requirements, 224–225 natural diet composition, 221–222 species differences in requirements, 208–209, 220–221 marine fish broodstock requirements, 219–220 embryogenesis and early larval requirements, 211–213 food fish composition, 207 juvenile and adult requirements, 209, 218–219 larval and early postlarval requirements, 208, 213–218 plankton composition, 207, 210 species differences in requirements, 208–209 oxidation ATP yield, 762 odd-chain fatty acids, 762 overview, 760, 762 peroxidation, see Fatty acid peroxidation prospects for aquaculture nutrition, 244–246 psychiatric disorder nutritional therapies, 200 special feeds n-3 highly unsaturated fatty acids, 658–661 n-6/n-3 ratio, 661 sources, 659–661 structures highly unsaturated fatty acids, 206–207 monounsaturated fatty acids, 189 nomenclature, 187–188 polyunsaturated fatty acids, 189, 206 saturated fatty acids, 188 Fatty acid peroxidation consequences, 233–236 mechanisms, 232–233, 535 oxidized lipid feeding effects, 462, 464, 621–622 protectants ascorbic acid, 238–239, 622 carotenoids, 238 feed additives, 536 synergistic antioxidants, 238–239 tocopherols, 236–238, 622 813 FE, see Fecal energy Feather meal, protein supplementation, 518 Fecal energy (FE), nutritional energetics, 8–9 Fecal sample, collection from fish, 9–10 Feed conversion ratio, feed evaluation, 589 Feed formulation and manufacture, see Diet formulation and manufacture Feeding larva, see Larva level and frequency effects on digestibility, 16–17 regimens, see also individual fish species deprivation and disease resistance, 694–695 medicated feeds, 697–698 nutritional condition manipulation, 695–696 seasonal feeding, 696–697 stimulants, 388–389 Fermentation, heat loss, 38 Fiber feed composition, 536–537 proximate analysis, 580–581 role in fish nutrition, 464 Fingerling feed, characteristics, 543 Fish diet disease, historical perspective of study, 63 Fish meal animal feed and fertilizer production, 665, 667 manufacturing process, 662–663 protein supplementation, 519 supply, 654, 661 total utilization of fish, 663 Fish oil, quality standards for salmonid diets, 524 Fish protein concentrate (FPC) hydrolyzed fish protein production, 664–665 production, 663 FMN, 73 Folic acid antimetabolites and inactivation, 93–94 clinical assessment of status, 94–95 deficiency syndrome, 91–92, 476 discovery, 89 functions, 90–91 immune system effects, 686 requirements, 92 sources and preparations, 92–93 structure and physical properties, 89–90 Formulation of diets, see Diet formulation and manufacture FPC, see Fish protein concentrate 814 Index Fry fatty acid requirements, 224–225 feeds, 541–542 Fumonisin B1 , manifestations in fish, 629–630 G Gallbladder anatomy, 411–412 function, 412–413 Gastrin, characteristics in digestion, 430 Gastrin-releasing peptide, characteristics in digestion, 430 Gastroenteropancreatic hormones, digestion control and regulation, 428–431 GE, see Gross energy Gelatin, protein supplementation, 519 Genera, aquaculture species, 804–805 Gill ammonia elimination, 349, 757, 765 raker structure and function, 392 Gizzard, function, 406, 415, 418 Glucagon glycogen metabolism regulation, 321 receptor and signaling, 323 β-Glucan immunostimulation, 530, 690–693 supplementation, 530 Glucocorticoids, glycogen metabolism regulation, 321 Glucose blood glucose regulation, 324 gluconeogenesis, 320, 759 glycolysis, 316, 757–759 nutritional energetics, 7, 40 pentose phosphate pathway, 317–318, 759 Glucosinolates sources, 608–610 thyroid effects in fish, 609–611 Glutamate, transport, 764 γ -Glutamyltransferase, nutrient transport role, 426 Glycogen metabolism blood glucose regulation, 324 gluconeogenesis, 320, 759 glucosidic pathways of breakdown, 324 glycogenolysis, 759–760 glycolysis, 316, 757–759 hormonal control, 320–323 Krebs cycle, 318–319 muscle, 327 oxidative phosphorylation, 319–320 pentose phosphate pathway, 317–318, 759 synthesis, 320 Glycogen phosphorylase, regulation, 322 Glycogen synthase, regulation, 322–323 Glycolysis ATP yield, 316, 757–759 overview, 316, 757–759 Gonads, energy content, 47–48 Gossypol cottonseed source and inactivation, 614–615 manifestations in fish, 615–616 Grinding, feed manufacture, 558–560 Gross energy (GE), nutritional energetics, Grower feed, characteristics, 543 Growth hormone, supplementation, 532 Guelph Test Diet, specifications, 513 Gustation detectable molecules, 382, 387–388 feeding stimulants, 388–389 free amino acid pools, 346 neuroanatomy, 387 sensitivity, 384, 387 taste bud features, 384, 387 Gut microflora cellulase activity, 415 manipulation in aquaculture, 416 G w , see Daily instantaneous growth rate H H-440, specifications, 513 HE, see Heat production Heat increment of feeding (HiE) definition, 35 digestion and absorption processes, 37–38 factors affecting, 35–36, 42–43 metabolizable energy relationship, 36 recovered energy relationship, 36 species differences, 36 Heat loss, digestion and absorption processes enzymatic hydrolysis, 38 lipid absorption, 38 metabolic waste formation and excretion, 39 substrate transformation and retention, 39–43 Heat production (HE) calorimetric measurement direct calorimetry, 25 indirect calorimetry, 25–27 overview, 24–25 comparative carcass analysis, 27–28, 49 heat increment of feeding, 24 HEf, see Fasting heat production HEm, see Maintenance energy requirement Hemagglutinins, features and toxicity, 606–607 Index Hemapoiesis, clinical assessment of nutriton, 132 HFP, see Hydrolyzed fish protein HiE, see Heat increment of feeding Histamine, feed contamination, 621 Histidine, nutritional requirements by fish species, 156 Hitra disease, pathogenesis, 464 Hydrolyzed fish protein (HFP) production, 664–665 substitution for fish meal in feeds, 667 I IAAI, see Indispensible amino acid index Ideal protein concept, amino acid requirement estimation, 173–174 IFV, see International Feed Vocabulary Immune system components in fish, 674–675 nutrient effects arginine, 677, 682 ascorbic acid, 678–680, 684–686 carbohydrate levels in diet, 682 fatty acids n-3 fatty acids, 683–684 oxidized fatty acid, 682–683 folic acid, 686 iron, 681, 688 magnesium, 681, 687 manganese, 688 phosphorous, 681, 687 protein levels in diet, 677 pyridoxine, 687 selenium, 681, 687–688 vitamin A, 680, 686 vitamin E, 680, 686 zinc, 681, 688–689 prospects for study of nutritional interactions, 698–699 stimulants, see Immunostimulant Immunostimulant definition, 689 β-glucans, 530, 690–693 levamisole, 692–693 nonspecific immune response stimulants, 690–693 nucleotides, 692–693 specific immune response stimulants, 689–690 Indispensible amino acid index (IAAI), protein quality assessment in feeds, 588 Innervation, digestive system, 429 Inositol antimetabolites and inactivation, 109–110 clinical assessment of status, 110 815 deficiency effects, 109, 474 discovery, 107–108 functions, 108–109 larva requirements, 227–231 requirements, 109 sources and preparations, 109 structure and physical properties, 108 Insect meal, protein supplementation, 522 Insulin glycogen metabolism regulation, 321–322 supplementation, 532 Intermediary metabolism, see also Carbohydrate; Lipid; Protein control mechanisms, 311–313 dietary effects, 765–766 organization, 310–311 prospects for study, 358–359 International Feed Vocabulary (IFV), classifications, 516 Intestinal bulb, distinguishing from true stomach, 403–404 Intestine anatomy, 403 histology, 403 ion exchange, 426 segments, 403 Iodine deficiency, 295–296, 481 functions, 295 metabolism, 295 requirement, 296 sources, 296 Iron deficiency, 287, 481 functions, 286–287 immune system effects, 681, 688 metabolism, 286–287 requirement, 288 sources, 288 toxicity, 481 Isoleucine, nutritional requirements by fish species, 157 Isotopes, nutritional energetics studies, 28 J Joule, caloric equivalents, K King, Charles Glen, contributions, 62, 99 Krebs cycle, overview, 318–319 Kries test, lipid quality testing in feeds, 586–587 Krill meal feed additive value, 656 protein supplementation, 521–522 816 Index L Larva artificial diet acceptability, 444–445 brine shrimp feeding of marine fish larvae, 214–215, 217, 229, 443–444 choline requirements, 227–231 eye development, 445–446 fatty acid requirements freshwater fish embryogenesis and early larva, 222–224 larva and fry, 224–225 marine fish embryogenesis and early larva, 211–213 larva and early postlarva, 208, 213–218 feed manufacture complex feeds, 577–578 microbound feeds crumbled cake feeds, 574–575 crumbled feeds, 573–574 flake feeds, 574 microencapsulated feeds, 576–577 on-size feeds microextrusion marumerization, 575–576 particle-assisted rotational agglomeration, 576 spray beadlets, 576 particle size, 573 first feeding development and timing, 432, 440–441 gut development embryogeness and digestion, 433 extracellular digestion, 440 fecal changes, 432–433 feeding before digestion development, 432 histology, 437, 439 protein assimilation efficiency, 439–440 striped bass development, 437, 439–440 inositol requirements, 227–231 lipid metabolism freshwater fish, 223–224, 229 marine fish, 211–213 ontogenic changes in life history strategies, 441–442 rotifer feeding of marine fish larvae, 214–215, 217 zooplankton feeding of marine fish larvae, 217–218, 442–444 Lavoiseier, Antoine, nutritional energetics contributions, 3–4 Lead, toxicity, 267, 484, 641 Least-cost feed analysis linear programming, 553–558 simultaneous equations, 551–553 Lectins, see Hemagglutinins Leucaena toxins, clinical manifestations, 494 Leucine, nutritional requirements by fish species, 158–159 Levamisole, immunostimulation, 691–693 Lipase, characteristics in digestion, 399–400, 421–422 Lipid, see also Fatty acid absorption, 421–422 analysis in feeds chemical tests of quality anisidine value, 586 hydrolytic rancidity, 585 Kries test, 586–587 oxidative rancidity, overview, 585–586 peroxide value, 586 Schall oven test, 587 thiobarbituric acid-reactive substances, 586 crude lipid proximate analysis, 580 biosynthesis, 193–194 classes and structures phosphoglycerides, 185–186 sphingolipids, 186 sterols, 186, 187 triacylglycerols, 184 wax esters, 184–185 deposition costs, 40–41 dietary levels antioxidant requirements, 205 correlation with tissue levels, 203–204 essential fatty acids, 204 fatty liver effects, 206 lipogenesis modification, 205–206 enzyme–substrate interactions, 182–183 feed ingredients, 523–525 heat loss of absorption, 38 high-energy diets, 202–203 lipases, 421–422 metabolism during embryogenesis and larval development freshwater fish, 223–224, 229 marine fish, 211–213 pellet top-dressing, 571 peroxidation, see Fatty acid peroxidation prospects for aquaculture nutrition, 244–246 recovered energy, 46–47 sources for farmed fish feeds fish oil alternatives, 239–243 marine fish larval feeds, 243–244 toxicity of oxidized lipids, 621–622 Index Lipoic acid functions, 113 structure and physical properties, 113 Liver amino acid uptake, 344 anatomy, 413 blood supply, 413 fatty liver, 206, 461–462 free amino acid pools, 347–349 functions, 427–428 glycogen vacuoles, 414 lipid storage, 414 nutrient processing, 428 Low-pollution feeds, characteristics, 544–547 Lysine arginine interactions, 160, 162 availability test, 584 dose–response, 351 nutritional requirements by fish species, 160–162 supplementation and metabolism, 351–352 utilization comparison between fish and other animals, 168–169 M Magnesium deficiency, 282–283 functions, 282 immune system effects, 681, 687 metabolism, 282 requirement, 283 sources, 283–284 Maintenance energy requirement (HEm) components, 33 measurement, 33–34 Manganese deficiency, 291, 482–483 functions, 290–291 immune system effects, 688 metabolism, 291 requirement, 291 sources, 291 Manufacture of diets, see Diet formulation and manufacture Maximum feed consumption (C max ), variability, 49 ME, see Metabolizable energy Meat meal, protein supplementation, 517–518 Megaloblastic anemia, nutritional deficiencies, 130–131 Mercury, toxicity, 267, 640 Mesotocin, receptor and signaling, 323 817 Metabolic fecal nitrogen (MFN), determination, 12–13 Metabolizable energy (ME) determination, 19–21, 40–41 efficiency of utilization, 39–40 factors affecting metabolic waste output dietary factors, 22–23 overview, 21–22 species differences, 23 land animal versus fish utilization, 707–708 lipid deposition costs, 40–41 protein deposition costs, 40–42 Methionine cysteine replacement values, 164 deficiency and cataracts, 355 detoxification functions, 356–357 nutritional requirements by fish species, 164–165 supplementation and metabolism, 355–357 MFN, see Metabolic fecal nitrogen Microextrusion marumerization (MUM), larval feed manufacture, 575–576 Milk by-products, protein supplementation, 518 Minerals, see also specific minerals bioavailability, 269–270, 480–481 bone metabolism, 264–265 deficiency signs, 277 distribution in fish, foods, and water, 263–264 dose–response, 262–263 essential elements, 260–261 metalloenzymes and functions, 261–262 osmoregulation and saltwater adaptation, 267–269 premixes, 526–527, 797–798, 803 prospects for aquaculture studies, 300–301 requirement study difficulties, 262–263 requirements, 279 supplementation effects in experimental diets, 270–271 synergism, 266 toxic elements and toxicity, 266–267, 484–485 transport mechanisms, 427 vitamin interactions, 265–266 Minimal metabolism, see Fasting heat production; Maintenance energy requirement Mirex, manifestations in fish, 635 Mitochondria, density and oxidative capacity in fish, 328–329 Mixing, feed manufacture, 560–562 818 Index Mouth structure and function, 389–391 teeth, 392 MUM, see Microextrusion marumerization Muscle carbohydrate metabolism adjustment of metabolic capacities growth, 331 species differences, 332 temperature, 329–330 fuel use during sustained activity, 327–329 glycogen regulation, 327 recuperation following exhaustive exercise, 326–327 regulation, 325–327 startle response, 325–326 fiber types and metabolic characteristics, 324–325 mitochondrial density and oxidative capacity, 328–329 stomach, 397 Mycotoxins, see also specific toxins aflatoxins, 489–490, 622–626 citrinin, 627–628 fumonisin B1 , 629–630 manifestations in fish, 489–490 ochratoxin A, 626 patulin, 626–627 rubratoxins, 627 sources, 489 T-2 toxin, 628 vomitoxin, 628 N NADPH, production and functions, 317–318 Nephrocalcinosis, pathogenesis, 486 Neuropeptide Y (NPY), characteristics in digestion, 431 NFE, see Nitrogen-free extract Niacin antimetabolites and inactivation, 86 clinical assessment of status, 86 deficiency syndrome, 84–85, 475 discovery, 83 functions, 84 requirements, 85 sources and preparations, 86 structure and physical properties, 83–84 Nitrogen-free extract (NFE), proximate analysis, 581 Nitrosamines, manifestations in fish, 639 NMR, see Nuclear magnetic resonance NPY, see Neuropeptide Y Nuclear magnetic resonance (NMR), nutritional energetics studies, 28 Nucleotides feeding stimulants, 388–389 immunostimulation, 692–693 Nutrient flow, 755,761 Nutrient retention , feed evaluation, 590 Nutritional disease, see also specific nutrients deficiency causes in farmed fish, 454–455 farmed versus wild fish, 454 specific dynamic action, 458 starvation, 455–456 stress theory, 457–458 O Ochratoxin A, manifestations in fish, 626 Olfaction detectable molecules, 382 epithelium, 384 feeding stimulants, 388–389 organ anatomy, 383–384 sensitivity, 384 special feed considerations, 656 OMP, see Oregon moist pellet Ontario Ministry of Natural Resources mineral premix, 527 MNR-98HS specifications, 512 vitamin premix, 526 Oregon moist pellet (OMP) bacterial contamination testing, 593 development, 509–510, 726 specifications, 510 Oregon Test Diet (OTD), specifications, 513 Orinthine–urea cycle (OUC), protein catabolism, 349–351 Osmoregulation, overview, 267–269 OTD, see Oregon Test Diet OUC, see Orinthine–urea cycle Oxidative phosphorylation ATP yield, 319–320 enzymes in electron transfer cascade, 763 nutritional energetics, P PAHs, see Polycyclic aromatic hydrocarbons Pancreas endocrine pancreas, 408, 429 exocrine pancreas, 407, 418 histology, 408, 411 location, 406 Pantothenic acid antimetabolites and inactivation, 82 clinical assessment of status, 82–83 deficiency syndrome, 81, 472, 474 discovery, 79–80 Index functions, 80–81 requirements, 81 sources and preparations, 81–82 structure, 80 PARA, see Particle-assisted rotational agglomeration Particle-assisted rotational agglomeration (PARA), larval feed manufacture, 576 Patulin, manifestations in fish, 626–627 PCBs, see Polychlorinated biphenyls Pellet binders materials, 527–528 pellet stability, 549 toxicity, 494–495 Pelleting cold pellet extrusion, 567 compressed pelleting, 563–565 cooling and drying, 569 crumbling and screening, 569–570 extruded dry pelleting, 565–566 mixtures and mix weights, 795–796 objectives, 563 shipping and storage, 571–572 shrimp feed, 750 top-dressing with lipids, 571 universal pellet cooker, 566–567 Penaeid shrimp amino acid requirements, 169 culture methods, 742–743 feeding behavior, 749–750 feeds and feeding natural foods, 748–749 practical feeds, 750 practices, 752–753 processing, 750–751 history of culture, 741 market, 741 nutrient requirements and sources, see also specific nutrients energy, 746 fatty acids, 747 minerals, 747–748 protein and amino acids, 149, 169, 745–746 vitamins, 747 protein requirements, 149 viral disease transmission from feed, 593 Pentose phosphate pathway, function, 317–318, 759 Pepsin characteristics in digestion, 398 hydrolyzed fish protein production, 664 protein digestibility test, 582–583 Pernicious anemia, nutritional deficiencies, 131 819 Peroxide value, 586 Pesticides, toxicity, 494 Phenylalanine nutritional requirements by fish species, 164–165 tyrosine replacement values, 165 Phloretin, urea excretion inhibition, 350 Phospholipase, characteristics in digestion, 400 Phosphorous absorption and transport, 275 bioavailability, 280–281, 481, 545–546 calcium interactions, 271–273 compounds, 274 deficiency, 276, 484 excretion, 281 fertilization of freshwater, 275–276 hormonal regulation, 273 immune system effects, 681, 687 low-pollution feeds, 544–547 output minimization in aquaculture, 281–282 requirements, 278 sources natural, 274–275 supplements, 278, 280 Photosensitizers, toxicity, 495, 497 Phytase, supplementation, 531 Phytic acid manifestations in fish, 607–608 mineral binding, 607 phosphorous bioavailability effects, 280–281, 481, 545–546 Phytoestrogens, fish effects, 608 Pigmentation, essential fatty acid role in marine fish, 216 Plant products adventitious toxins α-amylase inhibitor, 618–619 alkaloids, 612–614 cyclopropene fatty acids, 616–618 erucic acid, 611–612 glucosinolates, 608–611 gossypol, 614–616 hemagglutinins, 606–607 miscellaneous toxins, 619 overview, 603 phytic acid, 607–608 phytoestrogens, 608 soybean trypsin inhibitor, 603–606 protein supplementation, 520–521 starch from grains in feeds, 522–523 Polychlorinated biphenyls (PCBs) manifestations in fish, 635–638 uses, 635 820 Index Polycyclic aromatic hydrocarbons (PAHs), manifestations in fish, 638–639 Pond fish formula, specifications, 777 Pond fish vitamin premix, specifications, 778 Potassium deficiency, 284–285 functions, 284 metabolism, 284 requirement, 285 sources, 285–286 Poultry by-product meal, protein supplementation, 518 Probiotic administration, 531, 694 definition, 530, 693 mechanism of action, 693–694 safety, 694 Product quality feeds, characteristics, 547–548 Protein absorption, 419 amino acids, see Amino acids analysis in feeds biological tests biological value, 591 net protein utilization, 591 protein efficiency ratio, 591 protein retention, 591 chemical score, 587–588 chemical tests of quality biogenic amines, 584–585 lysine availability, 584 pepsin digestibility, 582–583 soybean meal tests, 583–584 total volatile nitrogen, 584 crude protein proximate analysis, 580 indispensible amino acid index, 588 composition of fish, 144 deficiency, 459–460 deposition amino acid requirements, 337 costs, 40–42, 335 estimation techniques, 334, 336–337, 766–767 feed intake effects, 767 fish size influences, 334–335 rates, 334, 766 tissue differences in synthesis, 766 end products of catabolism ammonia, 349–351, 764–765 urea, 349–351 energy utilization, 202 historical perspective of fish nutrition studies, 144 immune system effects of levels in diet, 677 land animal versus fish utilization, 707–708 proteases and peptidases, 419–421 recovered energy, 46–47 requirements factors affecting size and age, 149–150 water temperature, 150 gross requirements crustacea, 149 finfish, 145–148 maintenance requirements determination techniques, 150–151 estimates, 151 retention efficiency improvement, 768–769 sexual maturity effects on metabolism, 767 size effects on growth and protein synthesis, 338, 340 structure, 419–420 supplements animal by-products blood meal, 518 bone meal, 517–518 feather meal, 518 meat meal, 517–518 milk by-products, 518 poultry by-product meal, 518 classification by protein content, 517 fish products fish meal, 519 wet fish products, 519–520 gelatin, 519 insect meal, 522 krill meal, 521–522 plant products, 520–521 turnover, 144, 333 Proximate analysis ash, 581 crude fiber, 580–581 crude lipid, 580 crude protein, 580 ingredient categories, 578–579 nitrogen-free extract, 581 water, 579 Pyloric ceca anatomy, 401 brush border, 401, 403 function, 401, 403 Pyloric sphincter digestive enzyme characteristics, 398–400 function, 397, 401 Pyridoxine antimetabolites and inactivation, 79 clinical assessment of status, deficiency syndrome, 77–78, 472 Index discovery, 76 functions, 77 immune system effects, 687 requirements, 78 sources and preparations, 78–79 structure, 76–77 R RE, see Recovered energy Recirculation system feeds, characteristics, 547 Recovered energy (RE) definition, 43 estimation, 43 feed restriction effects, 45–46 heat increment of feeding relationship, 36 protein versus lipid deposition, 46–47 tissue type dependence, 43–44 trout composition studies, 44–45 Reproduction fatty acid oxidation as energy source, 196 nutritional energetics, 47–48 Respiratory quotient (RQ), determination, 26 Riboflavin antimetabolites and inactivation, 75 clinical assessment of status, 75–76 deficiency syndrome, 74, 472 discovery, 73 functions, 73–74, 472 requirements, 74–75 sources and preparations, 75 structure, 73 Romet, catfish treatment, 716 Rotifer, feeding of marine fish larvae, 214–215, 217 RQ, see Respiratory quotient Rubner, M., nutritional energetics contributions, 4, 38 Rubratoxins, manifestations in fish, 627 S Salmon culture methods, 721–722 fatty acid composition, 729 feeding practices automatic feeding, 731 frequency of feeding, 731–732 hand-feeding, 730 particle sizes of feeds, 731 flavor and color optimization of product, 729 formulation of feeds high-protein dry diet, 773 model diets, 727–728, 794 821 moist pellet formula and specifications, 772 pellets, 726 pollution minimization, 728–730 history of culture, 720 market, 720–721 nutrient requirements, see also specific nutrients energy, 724 fatty acids, 724 minerals, 725 protein and amino acids, 723–724 vitamins, 724–725 temperature and recommended feeding amounts, 793 Saltwater adaptation, overview, 267–269 Scale calcium metabolism, 272–274 structure, 265, 274 Schall oven test, lipid quality testing in feeds, 587 Scientific names, aquaculture species, 804–805 SCP, see Single-cell protein Screamer disease, pathogenesis, 480 SDA, see Specific dynamic action Seawater transfer feeds, characteristics, 547 Secretin, characteristics in digestion, 431 Sekoke disease, features, 497 Selenium deficiency and fatty acid effects, 237, 297–298 fatty acid peroxidation protection, 239 functions, 296–297 immune system effects, 681, 687–688 interactions with other nutrients, 266, 485 metabolism, 297 requirement, 298 sources, 298 Semipurified diets, manufacture, 572–573 Senecio alkaloids, toxicity, 492, 494 Serotonin, dietary trytophan and synthesis, 357–358 Sex reversal, hormone supplement manipulation, 532 SGR, see Specific growth rate Shrimp, see Brine shrimp; Penaeid shrimp Single-cell protein (SCP), lesions, 500 Sodium deficiency, 284–285 functions, 284 metabolism, 284 requirement, 285 sources, 285–286 822 Index Sodium/potassium-ATPase, saltwater adaptation role, 269 Somatostatin, characteristics in digestion, 431 Soybean meal quality tests, 583–584 utilization efficiency, 604–605 Soybean trypsin inhibitor heat inactivation, 604, 606 manifestations in fish, 603, 605–606 species sensitivity, 605–606 structure, 603 Spawning, energetics, 48 Special feeds definition, 652 fatty acids n-3 highly unsaturated fatty acids, 658–661 n-6/n-3 ratio, 661 sources, 659–661 formulation and manufacture factors aquaculture environment, 653–654 color and contrast, 656 digestibility, 656–657 nutritional components, 653 odor, 656 particle density, 655–656 particle texture, size, and shape, 657 physical and chemical properties, 654–655 species, 652–653 storage stability, 657–658 water stability, 655 manufacturing challenges, 661–665, 667–668 Specific dynamic action (SDA), feeding cost, 458 Specific growth rate (SGR), calculation, 50 Starvation, manifestations, 455–456 Stomach agastric fish nutrition, 416–417 compartments, 396 configurations, 393, 396 digestive enzyme characteristics, 398–400 function, 397 gastric glands, 396–397, 415 mucosal cells, 397 muscle, 397 tissue layers, 396 Stress environmental conditions affecting fish health, 673–674 nutritional disease, 457–458 Sulfonamides, toxicity, 500 Swimming, heat losses associated with activity, 34–35 T T-2 toxin, manifestations in fish, 628 Taste, see Gustation TBARS, see Thiobarbituric acid-reactive substances Teeth, structure and function, 392 Terramycin, catfish treatment, 716 Testosterone, supplementation, 532 Tetrodotoxin fish meal contamination, 619 structure, 620 TGC, see Thermal-unit growth coefficient Thermal-unit growth coefficient (TGC), calculation, 50–51 Thiamin antimetabolites and inactivation, 72 clinical assessment of status, 72 deficiency syndrome, 68, 471–472 discovery, 66 functions, 67–68 requirements, 68, 70–71 sources and preparations, 71–72 structure, 67 Thiobarbituric acid-reactive substances (TBARS), lipid quality testing in feeds, 586 Threonine, nutritional requirements by fish species, 166–167 Thyroid hormones glucosinolate effects, 609–611 mirex effects, 635 supplementation, 531–532 Tilapias culture practices, 733–735 digestibility coefficients of feedstuffs, 736 feeds and feeding natural foods, 737–739 practical feeds, 739–741 genera, 732 nutrient requirements, see also specific nutrients energy, 736 fatty acids, 736–737 minerals, 737 protein and amino acids, 735–736 vitamins, 737 temperature effects, 732 Total volatile nitrogen (TVN), protein quality test, 584 Toxaphene manifestations in fish, 632–633 uses, 632 Index Transition feed, characteristics, 542–543 Trout culture methods, 721–723 fatty acid composition, 729 feeding practices automatic feeding, 731 frequency of feeding, 731–732 hand-feeding, 730 particle sizes of feeds, 731 flavor and color optimization of product, 729 formulation of feeds model diets, 727–728, 794 pellets, 725–726 pollution minimization, 728–730 PR-6 specifications, 776 SD7 specifications, 774 history of culture, 720 market, 720–721 nutrient requirements, see also specific nutrients energy, 724 fatty acids, 724 minerals, 725 protein and amino acids, 723–724 vitamins, 724–725 temperature and recommended feeding amounts, 792 True digestible energy, determination, 11 Trypsin, characteristics in digestion, 398 Tryptophan brain uptake, 358 nutritional requirements by fish species, 167–168, 357 supplementation and metabolism, 357–358 TVN, see Total volatile nitrogen Tyrosine, replacement values for phenylalanine, 165 U UE, see Urinary energy loss Universal pellet cooker (UPC), pelleting, 566–567 UPC, see Universal pellet cooker Urea excretion, 350–351, 764 protein catabolism production, 349–351 Urinary energy loss (UE) excretion rates, 19–20 factors affecting metabolic waste output dietary factors, 22–23 overview, 21–22 species differences, 23 823 measurement, 20–21 sources, 18–19 Urolithiasis, histology, 486, 488 V Valine, nutritional requirements by fish species, 159 Vasoactive intestinal peptide (VIP), characteristics in digestion, 431 Vasotocin, receptor and signaling, 323 Vegetable oils, sources for farmed fish feeds, 239–243 VIP, see Vasoactive intestinal peptide Visceral granuloma, features, 485–486 Vitamin, see also specific vitamins animal protein factors, 128–129 avitaminosis, 63 cell permeability factors, 129–130 coenzyme activation factors, 130 deficiency syndromes, 69, 130–131 fat-soluble vitamins, overview, 113–114 historical perspective of study, 62–63 hypervitaminosis, 63 mineral interactions, 265–266 premixes, 525–526, 775, 778, 797–803 requirements by fish species, 70 test diets and conditions, 63–66 water-soluble vitamins, overview, 66 Vitamin A clinical assessment of status, 117–118 deficiency and excess syndromes, 116–117, 465 discovery, 114 fatty acid peroxidation protection by carotenoids, 238 functions, 116 immune system effects, 680, 686 requirements, 117 sources and preparations, 117 structure and physical properties, 114–115 Vitamin B12 antimetabolites and inactivation, 98 clinical assessment of status, 98–99 deficiency syndrome, 97 discovery, 95 functions, 96–97 requirements, 97–98 sources and preparations, 98 structure and physical properties, 95–96 Vitamin C, see Ascorbic acid Vitamin D clinical assessment of status, 120 deficiency and excess syndromes, 119, 466 discovery, 118 functions, 119 824 Index requirements, 120 sources and preparations, 120 structure and physical properties, 118–119 Vitamin E clinical assessment of status, 124 deficiency and excess syndromes, 123, 466, 469, 471 discovery, 120 fatty acid peroxidation protection synergistic antioxidants, 238–239, 622 tocopherols, 236–238, 622 food storage utilization, 237–238 functions, 122–123, 466 immune system effects, 680, 686 lipids in diet and antioxidant requirements, 205, 237 oxidation, 683 requirements, 123, 236, 466 sources and preparations, 123–124 structure and physical properties, 121–122 Vitamin K clinical assessment of status, 128 deficiency and excess syndromes, 126–127, 471 discovery, 124 functions, 125–126 requirements, 127 sources and preparations, 127–128 structure and physical properties, 125 Vitamin P, features, 129 Vomitoxin, manifestations in fish, 628 W Warfarin, vitamin K antagonism, 127–128 Water, proximate analysis, 579 Water activity, antimicrobial control in feeds, 534 Water quality, environmental conditions affecting fish health, 673–674 Water temperature digestibility influences, 17–18 feeding amount charts by temperature salmon, 793 trout, 792 metabolic regulation, 756 muscle adjustment of metabolic capacities, 329–330 protein requirement influences, 150 Wound healing, ascorbic acid role, 477–478 Z ZE, see Branchial energy loss Zinc availability in feeds, 539 deficiency, 293–294, 483–484 functions, 292 immune system effects, 681, 688–689 metabolism, 292–293 requirement, 294 sources, 294 Zooplankton fatty acid composition, 207, 210 feeding of marine fish larvae, 217–218, 442–444 nutrient composition, 442–443 ...Third Edition Fish Nutrition This Page Intentionally Left Blank Third Edition Fish Nutrition Edited by John E Halver School of Aquatic and Fishery Sciences University of Washington... Affecting Fish Health Dietary Components Influencing Fish Health Feeding Practices Affecting Fish Health Concluding Remarks and Research Needs References 672 673 675 694 698 699 Diet and Fish Husbandry... Dominique P Bureau (1), Fish Nutrition Research Laboratory, Department of Animal and Poultry Science, University of Guelph, Guelph, Ontario N1G 2W1, Canada C Young Cho (1), Fish Nutrition Research
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