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Mineral Nutrition Prelims 17/8/99 11:21 am Page i THE MINERAL NUTRITION OF LIVESTOCK 3rd Edition Mineral Nutrition Prelims 17/8/99 11:21 am Page ii Mineral Nutrition Prelims 17/8/99 11:21 am Page iii THE MINERAL NUTRITION OF LIVESTOCK 3rd Edition E.J. UNDERWOOD (deceased) and N.F. SUTTLE Moredun Research Institute Pentland Science Park Bush loan Penicuik Midlothian EH26 0PZ UK CABI Publishing Mineral Nutrition Prelims 17/8/99 11:21 am Page iv CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 10 E. 40th Street Wallingford Suite 3203 Oxon OX10 8DE New York, NY 10016 UK USA Tel: +44 (0)1491 832111 Tel: +1 212 481 7018 Fax: +44 (0)1491 833508 Fax: +1 212 686 7993 Email: [email protected] Email: [email protected] © CAB International 1999. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Underwood, Eric J. (Eric John), 1905– The mineral nutrition of livestock / E.J. Underwood and N.F. Suttle. -- 3rd ed. p. cm. Includes bibliographical references (p. ) and index. ISBN 0-85199-128-9 1. Minerals in animal nutrition. I. Suttle, N. F. II. Title. SF98.M5U5 1999 636.089527--dc21 99–11802 CIP ISBN 0 85199 128 9 Typeset in Garamond by Columns Design Ltd, Reading. Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn. Mineral Nutrition Prelims 17/8/99 11:21 am Page v Contents Preface vii 1. General Introduction 1 2. Natural Sources of Minerals 17 3. The Detection and Correction of Mineral Imbalances in Animals 47 4. Calcium 67 5. Phosphorus 105 6. Magnesium 149 7. Sodium and Chlorine 185 8. Potassium 213 9. Sulphur 231 10. Cobalt 251 11. Copper 283 12. Iodine 343 13. Iron 375 14. Manganese 397 15. Selenium 421 16. Zinc 477 v Mineral Nutrition Prelims 17/8/99 11:21 am Page vi vi Contents 17. Occasionally Beneficial Elements (Boron, Chromium, Lithium, Molybdenum, Nickel, Silicon, Tin, Vanadium) 513 18. Essentially Toxic Elements (Aluminium, Arsenic, Cadmium, Fluorine, Lead, Mercury) 543 19. Design of Supplementation Trials for Assessing Mineral Deprivation 587 Appendices 599 Index 601 Mineral Nutrition Prelims 17/8/99 11:21 am Page vii Preface It was with some trepidation that, some 7 years ago, I accepted an invitation from CAB International to revise the late Eric Underwood’s 1980 text for The Mineral Nutrition of Livestock. As a ‘raw’ post doctorate I had reviewed the third (1971) edition of his other major publication Trace Elements in Human and Animal Nutrition for CAB International and remember likening him to Goldsmiths ‘Village Teacher’ – ‘that one small head could carry all he knew’! I also had the privilege of hearing him sum up the proceedings of the first two international symposia on Trace Element Metabolism in Animals (TEMA 1 and 2) in Aberdeen (1968) and Wisconsin (1971): his ability to quickly distil the packed and complex proceedings of such gatherings and present the highlights with a touch of humour were truly remarkable. However, trace elements were only half of his repertoire and this book gathered together the essentials of macro- as well as trace element nutrition while focusing entirely on the practical needs of animals. My task has been simplified by the excellent foundation that Eric laid, the advances in information technology which now expedite any literature search and – sadly – a dramatic reduction in funding for applied research relating to animal nutrition in most developed countries, which took place towards the end of the last decade. Thus, the series of TEMA symposia, now approaching their 10th anniversary, were soon renamed Trace Elements in Man and Animals and the ‘Animal’ contribution has shrunk out of all recognition: this despite the fact that natural mineral imbalances frequently afflict animals and animal products are still important sources of minerals for man. The aim of the book must therefore, remain unchanged from that of the second edition ‘meeting the needs of under- graduate and graduate students of nutrition in colleges of agricultural science, animal husbandry and veterinary science, of teachers and research workers in animal nutrition, of agricultural and veterinary extension officers in developed as well as developing regions of the world and of progressive live- stock producers, wherever situated, who wish to apply modern scientific knowledge of mineral nutrition to their own enterprises’. vii Mineral Nutrition Prelims 17/8/99 11:21 am Page viii viii Preface It was also with trepidation that I made increasingly bold changes to the structure of Eric’s second edition, beginning by breaking the almost legendary links between calcium and phosphorus, and between sodium and potassium. There has been an explosion of interest in phosphorus, and the distinctive features of its metabolism which emerged merited a separate chapter: likewise, the contrasts between sodium and potassium become clearer when they are physically (though not physiologically) parted. Sulphur is now also given a chapter of its own, without demeaning the significance of its interactions with copper and molybdenum that were a feature of the second edition. The newer essential elements, boron, chromium, lithium, nickel, molybdenum, tin and vanadium have little practical nutritional significance, but this alone is worth emphasis and it is given under a new corporate heading of ‘Occasionally Beneficial Elements’. With fluorosis being an increasingly rare occurrence and other mineral toxicities causing increas- ing concern, coverage of fluorine is now given along with five other ‘Essentially Toxic Elements’, aluminium, arsenic, cadmium, mercury and lead in a single chapter. In the process of getting experts to critically review each draft chapter, I received the kind of comment that every author dreads: ‘of course I would not have structured the chapter that way’! This prompted a major rethink and a decision to depart from the original chapter structure which began with ‘Functions’, to one which begins with ‘Dietary sources’ of a given mineral and proceeds through ‘Metabolism’ to ‘Function’ and thence to the various aspects of dysfunction and its avoidance. Putting feed sources ‘up front’ showed just how sketchy the information was for most minerals, but it is better to reveal this than ignore it. Mineral imbalances are understood and controlled by exploiting sources and not prejudiced by skirting round numerous functions which never become rate-limited or health-limiting. Two recurring themes have been introduced to cope with the difficulties of assessing the likelihood of mineral-responsive disorders occurring on farms. One concerns the smooth, sequential patterns of biochemical change which usually accompany transition from a state of normality to one of dys- function and underlie the development of most disorders. Arising from this is the recognition that all biochemical measures of mineral status, from diet to urine, from erythrocyte to mitochondrion, from enzyme activity to spin- trapped electron, must be interpreted using a marginal band or ‘grey area’ which recognizes the imprecise link between what is measured and what is ‘felt’ functionally by the animal and financially by the farmer. The final revision was the addition of a concluding chapter on the role of ‘Supplementation Trials’ in resolving these continuing problems of defining whether or not mineral deficiencies have become sufficiently severe to impair performance and merit routine intervention. Since the first two editions were published, increasing pressure has been brought upon scientists to use international S.I. units of measurement when publishing results, i.e. a molar rather than mass basis. Compliance has varied from country to country due in part to opposing pressure from regulators of Mineral Nutrition Prelims 17/8/99 11:21 am Page ix Preface ix feed composition for animals and man who generally set limits in terms of mass. A compromise has been struck in this revision by using mass as the basis for reporting the mineral composition of feeds and for those minerals in animal tissues which are noted for the risks they might pose to man (e.g. arsenic, lead and cadmium): S.I. units are generally used for the assessment of mineral status of animal tissues and fluids. Whichever system is used, a conversion factor is usually close at hand. Comments and suggestions for inclusion or omission in any subsequent revision can be sent by e-mail to [email protected] or [email protected]. I am indebted to the Moredun Foundation for Animal Health and Welfare for financial support and library facilities without which the revision would not have been possible. I am also grateful to the following colleagues for scrutinizing particular chapters: Dennis Scott (Rowett Research Institute (RRI), Aberdeen), Colin Whitehead (Roslin Institute, Roslin) and Ronald Horst (Iowa State University, Iowa) on calcium and phosphorus; Andrew Sykes (Lincoln University, New Zealand) on magnesium and cobalt; Clive Phillips (School of Veterinary Medicine, Cambridge) and Paul Chiy (University of Wales, Bangor) on sodium, potassium and chlorine; Bob Orskov (RRI) on sulphur; Alan MacPherson (Scottish Agricultural Colleges, Auchincruive) on copper; Ian Bremner (RRI) on iron and cadmium; John Arthur (RRI) on iodine and sele- nium; Roger Sunde (University of Columbia, Missouri) and Doug Jones (Moredun Research Institute, Penicuik) on selenium; and John Chesters (RRI) on zinc. Without their vigilance and advice, this revision would have been sadly lacking. I am even more indebted to my typist-cum-research assistant, Jocelyn Brebner, for her unbelievable patience and cheerfulness through seemingly endless redrafts. Finally, I must thank my equally patient wife, Tilly, who thought I had retired 3 years ago! N.F.