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V ~ gamin ~ ~ ' in erance of Animals Subcommittee on Vitamin Tolerance Committee on Animal Nutrition Boarc! on Agriculture National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1987

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National Academy Press 2101 Constitution Avenue, NW Washington, DC 20418 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 responsi- ble for the report were chosen for their special competences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The National Research Council was established 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 of advising the federal government. The Council operates in accordance with general policies determined by the Academy under the authority of its congressional charter of 1863, which establishes the Academy as a private, nonprofit, self-governing membership corporation. The Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in the conduct of their services to the government, the public, and the scientific and engineering communities. It is adminis- tered jointly by both Academies and the Institute of Medicine. The National Academy of Engineering and the Institute of Medicine were established in 1964 and 1970, respectively, under the charter of the National Academy of Sciences. This study was supported by the U.S. Department of Agriculture, Agricultural Research Service, under Agreement No. 59-32U4-5-6, and by the Center for Veterinary Medicine, Food and Drug Administration of the U.S. Department of Health and Human Services, under Cooperative Agreement No. FD-U-000006-06-1. Additional support was provided by the American Feed Industry Association, Inc. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authoring subcommittee and do not necessarily reflect the views of the sponsors. Library of Congress Cataloging-in-Publication Data National Research Council (U.S.). Subcommittee on Vitamin Tolerance. Vitamin tolerance of animals. Bibliography: p. Includes index. 1. Vitamin tolerance in animals. I. Title. SF98.V5N37 1986 636.08'52 86-28466 ISBN 0-309-03728-X Copyright C) 1987 by the National Academy of Sciences No part of this book may be reproduced by any mechanical, photographic tic. or electronic process, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted, or otherwise copied for public or private use, without written permission from the publisher, except for the purposes of official use by the United States Government. Printed in the United States of America First Printing, January 1987 Second Printing January 1989 Third Printing,~ovember 1990

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Preface Optimal animal health and productivity are achieved by providing an animal with the correct amount and form of each essential nutrient. Good nutritional prac- tice requires supplementing practical diets with vitamin levels that exceed the bare minimum needed to prevent deficiency diseases. Yet, vitamins added to supplement feeds, which may have lost vitamins during processing and storage, or to protect the animal from stress or stim- ulate its immune system may necessitate the use of much higher levels. The goal in animal feeding is to supplement diets with vitamin levels that are adequate for nutritional needs and accommodate the practical conditions of feed manufacture and storage. To address these concerns, the Board on Agricul- ture's Committee on Animal Nutrition appointed the Subcommittee on Vitamin Tolerance in 1983. The sub- committee's report is the most comprehensive sum- mary of current data about the vitamin tolerances of animals. It will be helpful to livestock producers, live- stock extension specialists, animal nutritionists, animal nutrition students, and others interested in the subject. The presumed upper safe levels of vitamins in this book are meant to be used as guidelines to ensure that vitamin supplementation does not adversely affect ani- mal health. The subcommittee's objectives were to fo- cus on: (1) vitamin tolerance of domestic and laboratory animals under different nutritional and physiological states, (2) biological measures that can be used as crite- ria to establish tolerance, and (3) areas of incomplete knowledge. The first two objectives were, unfortunately, impos- sible to fulfill completely. For most of the vitamins, in- formation in the scientific literature on vitamin toxicities was incomplete with respect to the range of species studied and the quantitative aspects, such as dose-response definitions. In some cases, the literature indicated rather clearly the ranges of intake that would produce certain adverse responses in certain species; in others, it indicated little evidence of toxicity. The third objective is significant. The subcommit- tee's indicated values for the presumed safe levels are limited in scope by gaps in current information. Areas lacking adequate information are identified. It is the subcommittee's hope that this report points to the limits of current knowledge and important areas of research that can contribute to improvements in animal nutrition. The following individuals were responsible for re- spective sections of the report: Robert Blair, riboflavin, vitamin B6 (pyridoxine), biotin, vitamin Bit, and choline; Gerald F. Combs, fr., vitamin E and niacin; John W. Hilton, ascorbic acid and thiamin; Ronald L. Horst, vita- min D; George E. Mitchell, Tr., vitamin A; and John W. Suttie, vitamin K, pantothenic acid, and folic acid. The report was reviewed by the Committee on Ani- mal Nutrition, the Board on Agriculture, and outside reviewers. The subcommittee is grateful for the efforts of these individuals. We also thank Annette Bates, Es- ther Collins, Karen Davis, Andra Hinds, Frances New- some, and Pamela Senter, at our respective institutions, for their administrative assistance. The subcommittee especially acknowledges the contributions of Selma Baron, who served as staff officer during the early prep- aration of this report, and assistant editor Grace Jones Robbins, who helped us complete our task. GERALD F. COMBSS JR Chairman Subcommittee on Vitamin Tolerance -

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J SUBCOMMITTEE ON VITAMIN TOLERANCE GERALD F. COMBS, JR., Chairman, Cornell University ROBERT BLAIR, University of British Columbia JOHN W. HILTON, University of Guelph RONALD L. HORST, National Animal Disease Center, U.S. Department of Agriculture GEORGE E. MITCHELL, JR., University of Kentucky JOHN W. SUTTIE, University of Wisconsin, Madison .~. COMMITTEE ON ANIMAL NUTRITION JAMES G. MORRIS, Chairman, University of California-Davis FRANK AHERNE, University of Alberta RICHARD E. AUSTIC, Cornell University JIMMY H. CLARK, University of Illinois DONALD E. JOHNSON, Colorado State University ROY J. MARTIN, JR., University of Georgia FREDRIC N. OWENS, Oklahoma State University GARY L. RUMSEY, Tunison Laboratory of Fish Nutrition, U.S. Department of the Interior DALE R. WALDO, Animal Science Institute, U.S. Department of Agriculture Staff CARLA CARLSON, Reports Officer and Senior Editor GRACE NONES ROBBINS, Assistant Editor 1V

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BOARD ON AGRICULTURE WILLIAM L. BROWN, Chairman, Pioneer Hi-Bred International, Inc. JOHN A. PING, National Research Council Senior Fellow PERRY L. ADKISSON, Texas A&M University C. EUGENE ALLEN, University of Minnesota JOSEPH P. FONTENOT, Virginia Polytechnic Institute and State University ROBERT M. GOODMAN, Calgene, Inc. RALPH W. F. HARDY, Cornell University and BioTechnica International, Inc. ROGER L. MITCHELL, University of Missouri CHARLES C. MUSCOPLAT, Molecular Genetics, Inc. ELDOR A. PAUL, Michigan State University VERNON W. RUTTAN, University of Minnesota THOMAS D. TRAUTMAN, General Mills, Inc. JAMES G. TEER, Welder Wildlife Foundation JAN VAN SCHILFGAARDE, Agricultural Research Service, U.S. Department of Agriculture VIRGINIA WALBOT, Stanford University CONRAD I. WEISER, Oregon State University CHARLES M. BENBROOK, Executive Director v

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Contents INTRODUCTION VITAMIN A .... VITAMIN D .... VITAMIN E ....... VITAMIN K ..... ASCORBIC ACID THIAMIN NIACIN ..... RIBOFLAVIN (VITAMIN B2) VITAMIN B6 (PYRIDOXINE) Formic ACID PANTOTHENIC ACID . BIOTIN VITAMIN BI2 CHOLINE RESEARCH NEEDS . . . . SUMMARY ......... APPENDIX TABLE . . INDEX .11 .23 .31 .36 .53 . . . .64 .67 .70 .77 .85 .88 e V11 .93

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Figures and Tables FIGURES 1. Major compounds of the vitamin A group, 4 2. Chemical structures of vitamin D3 and vitamin D2, 12 3. Factors regulating 1,25-dihydroxyvitamin D and 24,25-dihydroxyvitamin D bio- synthesis, 13 4. Scheme for the pathogenesis of vitamin toxicosis, 14 5. Relationship between plasma and milk levels of vitamin D in the cow, 16 6. Relationship between plasma and milk levels of 25-OH-D in the cow, 16 7. Chemical structures of naturally occurring vitamin E-active compounds and ana- logues, 24 8. Chemical structures of three major forms of vitamin K, 32 9. The reduced and oxidized forms of ascorbic acid, 37 10. Chemical structure of thiamin hydrochloride, 44 11. Chemical structures of major niacin-active compounds, 48 12. Chemical structures of riboflavin and its coenzyme forms, 54 13. General chemical structure of vitamin B6 (pyridoxine), 59 14. Chemical structures of folic acid and 5-methyl-tetrahydrofolate, 65 15. Chemical structures of pantothenic acid and coenzyme A, 68 16. Chemical structure of biotin, 71 17. Chemical structure of the cyano form of vitamin By (cyanocobalamin), 76 18. Chemical structure of choline, 78 ~ vail

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Figures and Tables ix TABLES 1. Relative Vitamin A Activity of Carotenoids, 5 2. Research Findings of High Levels of Vitamin A in Animals, 6 3. Required and Presumed Upper Safe Levels of Vitamin A, 8 4. Vitamin D Sterols Used in Human and Animal Nutrition, 12 5. Research Findings of High Levels of Vitamin D in Animals, 17 6. Estimation of Safe Upper Dietary Levels of Vitamin D3 for Animals, 20 7. Relative Biopotencies of Vitamin E-Active Compounds and Analogues, 25 8. Research Findings of High Levels of Vitamin E in Animals, 26 9. Research Findings of High Levels of Vitamin K in Animals, 34 10. Research Findings of High Levels of Ascorbic Acid in Animals, 38 11. Research Findings of High Levels of Thiamin in Animals, 45 12. Research Findings of High Levels of Niacin in Animals, 50 13. Research Findings of High Levels of Riboflavin in Animals, 56 14. Research Findings of High Levels of Vitamin Be (Pyridoxine) in Animals, 60 15. Research Findings of High Levels of Biotin in Animals, 72 16. Research Findings of High Levels of Vitamin BE in Animals, 75 17. Research Findings of High Levels of Choline in Animals, 79 18. Appendix Table: Estimated Vitamin Requirements of Domestic and Laboratory Animals (Dry Diet Bases), 88

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