Digestive Physiology and Metabolism in Ruminants

Digestive Physiology and Metabolism in Ruminants Proceedings 0/ the 5th International Symposium on Ruminant Physiology, held at Clermont - Ferrand, on 3rd-7th September, 1979

Edited by

Y. Ruckebusch Head, Department (?t' Physiology, National Veterinary Sc/wol, Toulouse amI

P. Thivend Head, Laborotory of Ruminont Digestion, INRA, Theix

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MTP~R.~SS LIMITED

Imm/(/tiOlut! Medim[ nthlishers

Published by MTP Press Limited Falcon House Lancaster, England Copyright © 1980 MTP Press Limited Softcover reprint of the hardcover 1st edition 1980 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior permission from the publishers. British Library Cataloguing in Publication Data International Symposium on Ruminant Physiology, 5th Clermont - Ferrand, 1979 Digestive physiology and metabolism in ruminants. 1. Ruminants - Physiology - Congresses 2. Digestive organs - Mammals - Congresses 1. Title 1I. Ruckebusch, Yves III. Thivend, Pierre

599'.735'04132

QL737.U5

ISBN-13: 978-94-011-8069-6 e-ISBN-13: 978-94-011-8067-2 DOI: 10.1007/978-94-011-8067-2 REDWOOD BURN LlMITED

Trowbridge & Esher

Contents List

0/ Contributors

ix

Pre/ace

xiii

In appreciation 0/ A. T. Phillipson R. J. Moir

xv

Acknowledgements

xxv

Sponsors

xxvi

Historical profile of early digestive studie:;, Y. Ruckebusch SECTION I

GASTROINTESTINAL MOTILITY

2 The mixing and propulsion of the stomach contents of ruminants R. S. Wyburn

35

3 Rhythms of abomaso-intestinal motility L. Bueno and J. Fioramonti 4 The mechanisms controlling abomasal emptying and secretion F. R. Bell

53

SECTION"

81

BEHAVIOURAL PHYSIOLOGY AND NUTRITION

5 Ingestive behaviour and related activities in ruminants J. P. Dulphy, B. Remond and M. Theriez

103

6 Learning and associated factors in ruminant feeding behaviour L. R. Matthews and R. Kilgour

123

7 Hormones and metabolites in the control of food intake J. M. Forbes

145

8 Central control of water and salt intake in goats and sheep K. Olsson and M. J. McKinley

161

SECTION III

MICROBIAL ECOSYSTEM IN THE RUMEN

179

9 Microbial ecology of the rumen R. A. Prins and R. T. J. Clarke v

DIGESTIVE PHYSIOLOGY AND METABOLISM IN RUMINANTS

10 Factors affecting microbial growth yields in the reticulo-rumen

D. G. Harrison and A. B. McAllan 11 Adherent rumen bacteria - their role in the digestion of plant material, urea and epithelial cells K.-J. Cheng and J. W. Costerton 12 Mathematical modelling in analyses ofruminant digestive function: philosophy, methodology and application R. L. Baldwin and L. J. Koong SECTION IV

251

271

291

309 325

349

MINERAL METABOLlSM

18 Influence of minerals in rumen microbial digestion M. Durand and R. Kawashima 19 Dietary composition and the absorption of trace elements by ruminants I. Bremner and N. T. Davies 20 Calcium and phosphate homeostasis in ruminants and its relationship to the aetiology and prevention of parturient paresis A. D. Care, J. -Po Barlet and H. M. Abdel-Hafeez 21 Magnesium metabolism and hypomagnesaemia H. Martens and Y. Rayssiguier SECTION VI

227

RUMINANT DIGESTION AND ITS MANIPULATION

13 Digestion and end-product formation in the rumen from production rations J. D. Sutton 14 The effect of conservation and processing on the digestion of forages by ruminants D. J. Thomson and D. E. Beever 15 Possible nutritional constraints in meeting energy and amino acid requirements of the highly productive ruminant E. R. 0rskov 16 Chemical control of rumen microbial metabolism W. Chalupa 17 The pathophysiological effects of gastrointestinal and Iiver parasites in sheep J. D. Dargie SECTION V

205

375

409

429

447

INTERMEDIARY METABOLISM

22 Energy costs of digestion and metabolism in the gut

A. J. F. Webster vi

469

CONTENTS

23 Propionate metabolism and vitamin B 12 J. M. Elliot 24 Amino acid metabolism and hormonal control during growth A. H. Trenkle

485

25 Lactic acid metabolism D. Giesecke and M. Stangassinger

523

26 Mobilization, turnover and disposition of adipose tissue lipids R. S. Emery

541

SECTION VII

505

DIGESTIVE ADAPTATION

27 Digestive adaptation in the preruminant P. Thivend, R. Toullec and P. Guilloteau

561

28 Adaptation to changes in dietary composition, level and frequency of feeding W. Kaufmann, H. Hagemeister and G. Dirksen

587

29 Adaptation to diets containing significant amounts of non-protein nitrogen J. A. Nikolic, A. Pavlicevic, D. Zeremski and D. Negovanovic

603

30 Utilization of tropical feeds by ruminants T. R. Preston and R. A. Leng

621

31 Processing of animal waste by feeding to ruminants J. P. Fontenot and V. Jurubescu

641

SECTION VIII

COMPARATIVE DIGESTIVE PHYSIOLOGY

32 Evolution of microbial digestion in mammals I. D. Hume and A. C. I. Warner

665

33 Microbial digestion: rumen versus large intestine C. E. Stevens, R. A. Argenzio and E. T. Clemens

685

34 Coprophagy and related strategies for digesta utilization H. Hörnicke and G. Björnhag

707

35 The role of the ruminant's digestive tract as a water reservoir A. Shkolnik, E. Maltz and I. Choshniak

73 I

36 The digestive physiology of wild ruminants R. N. B. Kay, W. v. Engelhardt and R. G. White

743

37 The place of herbivores in the agricultural ecosystems W. Jarrige

763

Workshops

825

Index

841 vii

List of Contributors I. BREMNER Department of Nutritional Biochemistry, Rowett Research Institute, Bucksburn, Aberdeen, UK

H. M. ABDEL-HAFEEZ Department of Hygiene and Food Control Faculty of Veterinary Medicine, Assuit University, Assuit, Egypt

L. BUENO Physio-pathologie digestive, Ecole Nationale Veterinaire, Toulouse 31076, France

R. A. ARGENZIO National Animal Disease Center, Agricultural Research Service, United States Department of Agriculture, Ames, Iowa 50010, USA

A.D.CARE Department of Animal Physiology and Nutrition, University of Leeds, Leeds, UK

c. C. BALCH

Department of Feeding & Metabolism, National Institute for Research in Dairying, Shinfield, Reading, UK

R. J. BALDWIN Department of Animal Science, University of California, Davis, California, USA

W. CHALUPA Department of Clinical Studies, School of Veterinary Medicine, University of Pennsylvania, Kennet Square, Pennsylvania, USA

J.-P. BARLET Laboratoire des Maladies Metaboliques, C.R.Z.V. de Theix, 63110 Beaumont, France

K.-J. CHENG Animal Science Section, Agriculture Canada Research Station, Lethbridge, Alberta, Canada

D.E.BEEVER Department of Animal Nutrition and Production, Grassland Research Institute, Hurley, Maidenhead, UK

I. CHOSHNIAK Department of Zoology, University of Tel-Aviv, Tel-Aviv, Israel R. T. J. CLARKE Applied Biochemistry Division, Department of Scientific and Industrial Research, Palmerston-North, New Zealand

F.R.BELL Department of Medicine, Royal Veterinary College, University of London, London NWI OTU, UK

E. T. CLEMENS c/o Dr. S. N. Gershoff, Director, Nutrition Institute, Tufts University, Medford, Massachusetts 02153, USA

G.8JÖRNHAG Department of Animal Physiology, Agricultural College of Sweden, S-750 07 Uppsala 7, Sweden ix

DIGESTIVE PHYSIOLOGY AND METABOLISM IN RUMINANTS J. P. FONTENOT Department of Animal Science, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, USA

J. W. COSTERTON Department of Biology, University of Calgary, Calgary, Alberta, Canada J.D. DARGlE Department of Veterinary Physiology, Glasgow University Veterinary School, Glasgow, UK present address Joint FAO/IAEA Division of Atomic Energy in Food and Agriculture, Wagramerstrasse 5, P.O. Box 100, A-1400 Vienna, Austria

J. M. FORBES Department of Animal Physiology and Nutrition, University of Leeds, Leeds LS21 IJY, UK D. GIESECKE Institut für Physiologie, Physiologische Chemie und Ernährungsphysiologie, Universität München, 8000 München 22, Veterinärstrasse 13, Federal Republic of Germany

N. T. DAVIES Department of Nutritional Biochemistry, Rowett Research Institute, Bucksburn, Aberdeen, UK

P. GUILLOTEAU Station de Recherches Zootechniques INRA, Centre de Recherches de Rennes, 35042 Rennes, France

G. DIRKSEN 2 Medizinische Tierklinik Universität, Universität München, 8000 München 22, Veterinärstrasse 13, Federal Republic of Germany

H. HAGEMEISTER Department of Animal Nutrition, Federal Dairy Research Center, Kiel, Federal Republic of Germany

J. P. DULPHY Elevage des Ruminants, INRA de Theix, 63110 Beaumont, France

D. G. HARRISON Department of Agricultural Biochemistry, University of Newcastle-upon-Tyne, Newcastle-upon-Tyne, UK

M.DURAND Station de Recherches de Nutrition, CNRZ, Institut National de la Recherche Agronomique, Jouy-en-Josas 78350, France

F. A. HARRISON Department of Physiology, Agricultural Research Council Institute of Animal Physiology, Babraham, Cambridge, UK

J. M. ELLIOT Animal Science Department, Cornell University, Ithaca, New York, USA R.S.EMERY Department of Dairy Science, Michigan State University, East Lancing, Michigan, USA

H.HÖRNICKE Institut für Zoophysiologie, Universität Hohenheim, Garbenstrasse 30, 7000 Stuttgart 70, Federal Republic of Germany

W. v. ENGELHARDT Physiologisches Institut, Tierärztliche Hochschule, 3000 Hannover 1, Federal Republic of Germany

I. D.HUME Department of Biochemistry and Nutrition, University of New England, Armidale, New South Wales, Australia

J. FIORAMONTI Physio-pathologie digestive, Ecole Nationale Veterinaire, ToulOlise 31076, France

R. JARRIGE Station d'Elevage, INRA de Theix, 63110 Beaumont, France

x

LIST OF CONTRIBUTORS

L. R. MATTHEWS Department of Animal Behaviour, Ruakura Agricultural Research Centre, Hamilton, New Zealand

V. JURUBESCU Romania Ministry of Agriculture, Bucharest, Romania W.KAUFMANN Department of Animal Nutrition, Federal Dairy Research Center, Kiel, Federal Republic of Germany

R. J. MOIR Department of Animal Science and Production, University of Western Australia, Nedlands, Western Australia 6009, Australia

R. KA WASHIMA Department of Animal Science, Kyoto University, Kyoto, Japan 606

D. NEGOVANOVIC Department of Animal Husbandry, Institute for Scientific Investigation "Agroekonomik", Padinska Skela, Belgrade, Yugoslavia

R.N.B.KAY Physiology Department, Rowett Research Institute, Bucksburn, Aberdeen, UK R. KILGOUR Department of Animal Behaviour, Ruakura Agricultural Research Centre, Hamilton, New Zealand

J. A. NIKOLIC Department of Physiology and Radiobiology, Institute for the Application of Nuc1ear Energy in Agricuiture, Veterinary Medicine and Forestry, Zemun, Belgrade, Yugoslavia

L. J. KOONG Department of Animal Science, University of California, Davis, California, USA

K.OLSSON Department of Animal Physiology, Swedish University of Agricuitural Sciences, Uppsala 750 07, Sweden

R. A. LENG Department of Biochemistry and Nutrition, Faculty of Rural Science, University of New England, Armidale, New South Wales, Australia

E. R. q,RSKOV Department of Applied Nutrition, Rowett Research Institute, Bucksburn, Aberdeen, UK

A. B. McALLAN Department of Nutrition, National Institute for Research in Dairying, Shinfield, Reading, U K

A. PAVLlCEVIC Department of Physiology and Nutrition of Domestic Animals, Faculty of Agriculture, University of Belgrade, Zemun, Belgrade, Yugoslavia

M. J. McKINLEY Howard Florey Institute of Experimental Physiologyand Medicine, University of Melbourne, Parkville, Victoria 3052, Australia

T. R. PRESTON Centro Domincano de Investigacion Pecuaria con cana de Azucar, CEAGANA, Santo Domingo, Dominican Republic

E.MALTZ Department of Zoology, University of Tel-Aviv, Tel-Aviv, Israel H.MARTENS Physiologisches Institut, Tierarztliche Hochschule, 3000 Hannover I, Federal Republic of Germany

R. A. PRINS Laboratory of Animal Nutrition, Zootechnical Institute, Utrecht, The Netherlands xi

DIGESTIVE PHYSIOLOGY AND METABOLISM IN RUMINANTS Y. RA YSSIGUIER Laboratoire des Maladies Metaboliques, INRA de Theix, 63110 Beaumont, France

P.TIDVEND Laboratoire dc la Digestion des Ruminants INRA, Centre de Recherches Zootechniques et Veterinaires de Theix, 63110 Beaumont, France

B.REMOND Elevage des Ruminants, INRA de Theix, 63110 Beaumont, France

D. J. THOMSON Department of Animal Nutrition and Production, Grassland Research Institute, Hurley, Maidenhead, UK

Y.RUCKEBUSCH Ecole Nationale Veterinaire, 23 Chemin des CapelIes, 31076 Toulouse, France

R. TOULLEC Station de Recherches Zootechniques INRA, Centre de Recherches de Rennes, 35042 Rennes, France

A. SHKOLNIK Department of Zoology, University of Tel-Aviv, Tel-Aviv, Israel

A.H.TRENKLE Department of Animal Science, lowa State University, Ames, lowa 50011, USA

M. STANGASSINGER Institut für Physiologie, Physiologische Chemie und Ernährungsphysiologie, Universität München, 8000 München 22, Veterinärstrasse 13, Federal Republic of Germany

A. C. I. WARNER Division of Wildlife Research, C.S.I.R.O., Canberra, Australia

C.E.STEVENS Departments of Physiology, Biochemistry and Pharmacology, New York State College of Veterinary Medicine, Cornell University, Ithaca, New York 14853, USA

A. J. F. WEBSTER Department of Animal Husbandry, University of Bristol, Langford, Bristol, UK R. G. WHITE Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99701, USA

I. J. F. STOBO Department of Feeding & Metabolism, National Institute for Research in Dairying, Shinfield, Reading, UK

R. S. WYBURN SchooI of Veterinary Studies, Murdoch University, Perth, Western Australia 6153, AustraIia

J. D. SUTTON Department of Feeding & Metabolism, National Institute for Research in Dairying, Shinfield, Reading, UK

D. ZEREMSKI Department of PhysioIogy and Nutrition of Domestic Animals, Faculty of Agriculture, University of Belgrade, Zemun, Belgrade, Yugoslavia

M. THERIEZ Elevage des Ruminants, INRA de Theix, 63110 Beaumont, France

xii

Preface Two questions could not be avoided in the avant-propos of this book; (i) what is the importance to man of ruminant livestock, and (ii) what results of practical relevance in the growing mountain of scientific verbiage could be found in the Proceedings of this Symposium. Herbivores are an integral and critical part of the natural ecosystem which must be preserved because of their impact on human welfare. Wh at makes ruminants especially important to man is that they can thrive on fibrous forage and are thus the only viable enterprise over much of the earth's surface where crop growing is impracticable. They contribute a wide array of products in addition to 50000 000 tonnes ofmeat (1977) and represent a 'capital reserve' that can be drawn upon in times of emergency: milk for example (450000000 tonnes) can make the difference between subsistence and starvation. About 60% of the world's meat and 80 % of the milk are produced by one third of the world ruminant population in the developed regions and as much as 99 % of the power for agriculture is provided by the ruminant population in developing countries. For the next two decades, a probable increase by 30 % for .cattle and buffalo and more than 40 % for sheep and goats is expected by improving health, fertility, nutrition and genetic potential rather than feed resources. The importance of ruminant digestive physiology, is linked, by drawing on the capabilities of the rumen bacteria, to a greater use of crop residues including straw, sugar-cane refuse, sawdust and organic wastes such as a sewage sludge and animal manures. A viable system of food production requires an efficient transfer of energy and exchange of nutrit;:nts between soil, plant, animal and micro-organisms. This relationship can be ignored only at our peril, especially ifworld consumption ofruminant livestock production by AD 2000 is projected to be an increase by 80 % for milk, beef and veal and nearly 100% for mutton and lamb. A lactating cow on a diet of waste roughage and urea produces more protein than it consumes. Rumen efficiency might be enhanced by rumen stimulants and digestive aids. About 44 % of the live weight of cattle slaughtered for meat is inedible to man but is of high nutritional value as a protein concentrate for animal feeding. In this sense, better knowledge of how lower-energy plant materials could be processed through herbivores seems to be achallenge for man as a manipulator of the food cycle. We hope this book will serve to increase that knowledge. Y.R. and P.T. xiii

A. T. Phillipson 1911-1977

In appreciation of Andrew Tindal Phillipson MA, PhD (Cantab), MRCVS, Hon.DVSc (Copenhagen), Hon.DVM (Ghent), Hon.DVM and S (Edin) R. J. MOIR

Two generations of scientists and students of the digestive physiology, metabolism and nutrition of ruminants have known Andrew Tindal Phillipson as a pioneer and major contributor in these fields. In aseries of experiments he and a small group of colleagues thrust aside the confusion of the dogmas of the time to achieve a basic understanding of the processes in the rumen. Rapid development in ruminant research throughout the world followed and today delegates from 50 countries are represented at this the Fifth International Symposium on Ruminant Physiology. Phillipson was an active participant in the earlier symposia and was Chairman of the organizing committee of the third meeting held in Cambridge in 1969 and Editor ofthe proceedings. Phillipson's sudden death on the 10th of January 1977, at the age of 66, whilst returning to Cambridge from a meeting in London, deprives us of the continuing support of a valued and highly respected colleague. His academic background and development give some insight to his breadth of interest and scientific knowledge, apart from the intellectual and musical interests shared with his wife, Rachel, and three sons. He firstly read Agriculture in Cambridge and found particular interests in nutrition, physiology and genetics. Graduating in 1931 he then studied further physiology at University College, London, be fore commencing veterinary studies at the Royal Veterinary College. He qualified MRCVS in 1936 and remained briefly as house surgeon. His ability and research potential was recognized by the award of two post-graduate scholarships by the College and these took him back to Cambridge to read for his PhO in 1938. The Oirector of the Institute of PathoJogy made facilities avaiJabJe to him to study the motor activity of the pouches of the sheep's stomach. An intense young man, he received encouragement from the staff of that Institute, and his academic development and mastery of xv

DIGESTIVE PHYSIOLOGY AND METABOLlSM IN RUMINANTS

skills and experimentation may be gauged from the ftow of his publications ofthetime. lt is now just 40 years since PhiIlipson presented the first publications on his work. That he had al ready decided on the direction of his research is possibly indicated by the introduction to the major paper in 1939 when he stated: 'The study of the movements of the various compartments of the stornach is the obvious starting point when undertaking a programme of research into the alimentary physiology of sheep'. Apart from the intrinsic value of the work on motor activity of the stornach, it was cIear that he was a bold and able experimentalist, and could generate new techniques and integrate others into the task in hand. Furthermore he demonstrated that he could complete work, write about it cIearly and simply and publish it. lt is cIear that he early developed an appreciation of the continuous ftow aspects of the ruminant system. The mechanisms and measurements of these ftows were an important part of his thinking and work for the next 25 years, pointing the way to quantitation of digestive events. When the Agricultural Research Council established the Unit of Animal Physiology in 1941 in Cambridge under the direction of Sir Joseph Bancroft, PhiIlipson,just completing post-graduate studies, started with it. Phillipson, initiaIly together with Dr R. A. McAnnaIly, and later a number of others combined their developed talents, knowledge and skiIls to establish the modern bases ofruminant physiology and nutrition. The work ofthe previous 50 years had been unable to interpret the fate of the volatile fatty acids and the micro-organisms in the rumen producing them. PhiIlipson (1942) raised the question of the value of the odd and even carbon lower fatty acids, and whether the organisms were 'present in sufficient quantity to form an important end-product of fermentation', then together with McAnnaIly in a companion paper, offered the foIlowing definition: 'The fate of the volatile fatty acids appears to be that they are absorbed before they reach the abomasum. This concIusion rests upon the circumstantial evidence that they rapidly disappear from the rumen when introduced in quantities in excess of that already present, that they do not pass to the abomasum and that they are stable for 8 hours when incubated in vitro in rumen digesta. If the conception that absorption can occur readily in the rumen or omasum is accepted, then the fact that the volatile fatty acids are always present in the rumen is due, not so much to the fact that they are end-products of fermentation but to the fact that they are continuaIly being produced'. The possibility that glucose was a significant end-product of ceIlulose fermentation was ruled out by its rapid fermentation in the rumen and its absence from the abomasum; some may have been presented as bacterial starch. The nature of the ruminal epithelium which 'until recently (this) had been sufficient reason to deter curiosity regarding absorption from that organ' (1944) while a difficulty, could not prevent experimental examination of the xvi

IN APPRECIATION OF ANDREW TINDAL PHILLIPSON

new evidence. The several papers by Phillipson, McAnnally and Bancroft between 1942 and 1944 establishing absorption of fatty acids from the rumen, their use by tissues, and that the quantity was 'sufficient to supply an appreciable part of the energy requirement of the anima!' (1944) are now history. Thus the basis of Kellner's finding at the turn of the century that finely divided cellulose had the same fattening capacity as starch came nearer to understanding. In 1945 Phillipson gave an address which indicated the great advances of the previous 5years in fact and in understanding. The final paragraph of that review (1946) highlights his understanding. He wrote, 'The important question is this: how much protein and polysaccharide does the animal receive in microbial form? It is incorrect to assurne that the products of digestion in the rumen are wholly-free chemical substances, such as fatty acids, which can be absorbed and used directly, or that the animal lives entirely on the organisms that grow in the rumen. The ruminant lives not only on fatty acids and micro-organisms but also on any food, starch or protein that leaves the rumen unaffected by the action of micro-organisms. The final answer must depend on quantitative measurements of the amount ofthese materials that become available to the animai'. He continued to pursue this question at the Rowett Research Institute when he took up the Headship of the Physiology Department in 1947. The measurement of flow was difficult, but he had commenced work while still with the ARC on ways and means of measuring the flow of digesta from the stornach. His publications in 1948 and 1952 are most important ones, as they established the dimensions of this flow, and with a physiologist's sensitivity also established the principles and limitations of the procedures. In the next few years he worked with a number of young colleagues in quantitating various aspects of the flow from the abomasum particularly with respect to protein flow. Although he established the essential bases for quantitation of digesta flow, he did not participate to any extent in the development of modern marker technology. The Physiology Department of the Rowett Institute was very active under Phillipson and he continued his contributions in digestive physiology, microbiology, organic and inorganic nutrition, metabolism and pathology and the training and development of a considerable number of post-graduates and young scientists. In 1963 PhiIlipson returned to his 'horne base' in Cambridge as Professor of Veterinary Clinical Studies. The administration and development of his new clinical studies department, the heavy load of committees he headed or contributed to and the increasing financial and community pressures on Universities were heavy work to the painstaking scientist and his research and his publication outputs were severely reduced. Nevertheless his publication list is extensive, and his many papers and communications to learned and professional societies on original work are coupled with reviews, books and xvii

DIGESTIVE PHYSIOLOGY AND METABOLlSM IN RUMINANTS

chapters in books, occasional papers and lectures which, while highlighting his scientific contributions, indicate his capacity as an educator in his fields of work. Because of his great knowledge and his scholarly perception and logic he was widely sought as an examiner, as an adviser and in discussion. Not a good lecturer in formal situations, he was, in contrast, lively and vigorous in individual discussion. As his life's work demonstrates, Andrew Phillipson was a man of considerable purpose, drive and ambition. To most he appeared as a quiet, and indeed a most modest man not given to flamboyant displays: at times, because of the fierce intensity he applied for his work, particularly during the course of difficult experiments, he appeared distant, cold and unapproachable. He was, in fact, a cultured man of great individual charm and wit and a warm and generous friend. It is fitting that Andrew Phillipson and his outstanding contributions to ruminant digestive physiology be both remembered and honoured by this Symposium.

xviii

BIBLIOGRAPHY OF PROFESSOR A. T. PHILLIPSON Papers presenting original work Anaesthesia in sheep. A. T. Phillipson and S. F. Barnet!. Veto Rec., 1939,51, 869-872. Permanent stornach fistulae in ruminants. A. T. Phillipson and J. R. M. Innes. Q. J. Exp. Physiol. 1939,29,333-341. The movements of the pouches of the stornach of the sheep. A. T. Phillipson. Q. J. Exp. Physiol. 1939,29,395-415. The fluctuations of pH and organic acids in the rumen of the sheep. A. T. Phillipson. J. Exp. Biol. 1942, 19, 186-198. Studies on the fa te of carbohydrates in the rumen of the sheep. A. T. Phillipson and R. A. McAnally. J. Exp. Biol. 1942,19, 199-214. Physiological action of acetic acid in living tissue. J. Bareroft, R. A. McAnally and A. T. Phillipson. Nature. 1943, 151, 304. The absorption of sodium ortho-iodohippurate from the rumen of lambs. J. Bareroft, R. A. McAnally and A. 1. Phillipson. J. Exp. Biol. 1944,20, 132-134. Absorption of volatile acids from the alimentary tract of sheep and other species. J. Bareroft, R. A. McAnally and A. T. Phillipson. J. Exp. Biol. 1944, 20, 120-129 The mechanism of absorption from the rumen as exemplified by the behaviour of acetic, propionic and butyric acids. J. F. Danielli, M. W. S. Hitchcock, R. A. Marshall and A. T. Phillipson. J. Exp. Biol. 1945, 22, 75-84.

Volatile acid in the alimentary tract of the dog. A. T. Phillipson. J. Exp. Biol. 1947, 23, 346-349. Volatile acid in the portal blood of sheep. P. Schambye and A. T. Phillipson. Nature. 1949,164,1094. Experiments with dried bracken (Plnis aquilina). K. J. Carpenter, A. T. Phillipson and W. Thomson. Br. Veto J. 1950, 106, 292-308. The development of motor responses in the stornach of the foetal sheep. D. L. Duncan and A. T. Phillipson. J. Exp. Biol. 1951, 28, 32-40. A comparison of the mixtures of acetic, propionic and butyric acid in the Turnen and in the blood leaving the rumen. P. Kiddie, R. A. Marshall and A. T. Phillipson. J. Physiol. 1951,113,207-217. The absorption of acetate, propionate and butyrate from the rumen of sheep. M. J. Masson and Ä. T. Phillipson. J. Physiol. 1951, 113, 189-206. The passage of digesta from the abomasum of sheep. A. T. Phillipson. J. Physiol. 1952,116,84-97. The composition of the digesta leaving the abomasum of sheep. M. J. Masson and A. T. Phillipson. J. Physiol. 1952, 116, 98-111. The administration of cobalt by different routes to lambs maintained on a low-cobalt die!. A. T. Phillipson and R. L. Mitchel!. Br. J. Nutrit. 1952, 6, 176-189.

Volatile acid in the digesta ofruminants and other animals. S. R. Elsden, M. W. S. Hitchcock, R. A. Marshall and A. T. Phillipson. J. Exp. Biol. 1946,22,191.

The fatty acids present in the rumen of lambs fed on a flaked maize ration. A. T. Phillipson. Br. J. Nutrit. 1952,6, 190-198. XIX

DIGESTIVE PHYSIOLOGY AND METABOLlSM IN RUMINANTS The movement of potassium, sodium, chloride and water across the rumen epithelium of sheep. D. Parthasarathy and A. T. Phillipson. J. Physiol. 1953, 121,452-469. The rates of absorption of acetic, propionic and n-butyric acids. w. H. pfander and A. T. Phillipson. J. Physiol. 1952, 122, 102-110. Studies on the toxicity of bracken. A. T. Phillipson and R. S. Reid. J. Comp. Path. Ther. 1954, 64, 243-259. A histological study of the organisation of the rumen epithelium of sheep. M. J. Dobson, W. Brown, A. Dobson and A. T. Phillipson. Q. J. Exp. Physiol. 1956, 41, 247-253. Thiamine in the contents of the alimentary tract of sheep. A. T. Phillipson and R. S. Reid. Br. J. Nutr. 1957,11,27-41. Electrolyte concentrations in the erythrocytes of the goat and ox. J. y. Evans and A. T. Phillipson. J. Physiol. 1957,139,87-96. The absorption of chloride ions from the reticulo-rumen sac. A. Dobson and A. T. Phillipson. J. Physiol. 1958,140,94-104. Distension of the rumen and salivary secretion. A. T. Phillipson and C. S. W. Reid. Nature. 1958,181,1722-1723.

The assimilation of ammonia nitrogen by bacteria of the rumen of sheep. A. T. Phillipson, M. J. Dobson, T. H. Blackburn and M. Brown. Br. J. Nlitrit. 1962, 16, 151. Lipids of sheep lymph: Transport from the intestine. L. Felinski, G. A. Garton, A. K. Lough and A. T. Phillipson. Biochem. J. 1964, 90, 154-160. The absorption of calcium and magnesium from the rumen and small intestine of the sheep. A. T. Phillipson and J. E. Storry. J. Physiol. 1965, 181, 130-150. The influence of diet on the nitrogenous components passing to the duodenum and through the lower ileum of sheep. E. M. W. Clarke, G. E. Ellinger and A. T. Phillipson. Proc. R. Soc. B. 1966, 166,63-79. The flow of organic and inorganic materials through the alimentary tract of sheep. J. Bruce, E. D. Goodall, R. N. B. Kay, A. T. Phillipson and L. E. Vowles. Proc. R. Soc. B. 1966,166,46-62. Nitrogen metabolism in the ovine stornach : 1. The transfer of urea from the blood to the Turnen. c. J. F. Harrop and A. T. Phillipson. J. Agric. Sei. 1974,82(3),399-408. Original Contributions made to Societies Absorption of volatile acids from the rumen of sheep. R. A. McAnally and A. T. Phillipson. J. Physiol. 1942,101, 13P. Absorption of sodium ortho-iodo-hippurate from the rumen of lambs. R. A. McAnally and A. T. Phillipson. J. Physiol. 1942,101, 14P. The production of acetic, propionic and butyric acids in the alimentary canal. J. Barcroft, R. A. McAnally and A. T. Phillipson. Biochem. J. 1944,38, ii. Absorption of acetic, propionic and butyric acids from the alimentary tract. J. Barcroft, R. A. McAnally and A. r. Phillipson. Biochem. J. 1944,38, iii. The destruction of acetic acid by the beating hear!. J. Barcroft, R. A. McAnally and A. T. Phillipson. Biochem. J. 1944, 38. iv.

Assimilation of ammonia nitrogen by rumen bacteria. A. T. Phillipson, M. J. Dobson and T. H. Blackburn. Nature. 1959,183,402-404. Responses of the salivary glands to distension of the oesophagus and rumen. R. N. B. Kay and A. T. Phillipson. J. Physiol. 1959,148,507-523. Bloat in Cattle. XVI. Bovine saliva. The chemicaI composition of the parotid, submaxillary and residual secretions. A. T. Phillipson and J. L. Mangan. N.Z. J. Agric. Res. 1959,2,990-1001. The rate of transport of food along the digestive tract of sheep. J. P. Hogan and A. T. Phillipson. Br. J. Nutrit. 1960,14,147-155. The measurement of the flow of food to the duodenum of sheep. L. E. Harris and A. T. Phillipson. J. Anim. Prod. 1962,4,97-116.

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IN APPRECIATION OF ANDREW TINDAL PHILLIPSON The production and absorption of volatile acids in the rumen. R. A. Marshall and A. T. Phillipson. Proc. Nutr. Soc. 1945, 3, 238. The tolerance of sheep to low concentrations of blood sugar. M. W. S. Hitchcock and A. T. Phillipson. J. Physiol. 1946,105, 42P. The relation between the proportions of acetic and propionic acids passing from the rumen of sheep. R. A. Marshall and A. T. Phillipson. Proc. X VIf International Physiological Congress, Oxford. 1947, p. 358. A method of measuring the f10w of digesta from the stornach of sheep. A. T. Phillipson. J. Physiol. 1948,107, 21P. The passage of food through the stornach of sheep. A. T. Phillipson, R. Green, R. S. Reid and L. E. Vowles. Br. J. Nutrit. 1949,3, iii. The absorption of acetate from the rumen of sheep. A. T. Phillipson. J. Physiol. 1949, 109, I3P. Experimental cobalt deficiency. A. T. Phillipson and R. L. Mitchell. Br. J. Nutr. 1950,4, iii. The forces moving chloride ions through the rumen epithelium. A. Dobson and A. T. Phillipson. J. Physiol. 1954,125, 26-27P. The movements of ions across the reticulorumen sac. A. Dobson and A. T. Phillipson. Proc. XXth International Physiological Congress, Brussels, 1956,248-249. The influence of the contents of the rumen and of adrenalin upon its blood supply. A. Oobson and A. T. Phillipson. J. Physiol. 1956,133, 76-77P. The effects of distension of the oesophagus and rumen of sheep on salivation. R. N. B. Kay and A. T. Phillipson. J. Physiol. 1957,139, 7P. The incidence of pressure waves in the rumen of cattle. A. T. Phillipson and C. S. W. Reid. Proc. Nutr. Soc. 1960, 19, xxvii. The influence of urea and other dietary supplements on the nitrogen content of the digesta passing to the duodenum of hay-fed sheep.

R. N. B. Kay and A. T. Phillipson. Proc. Nutr. Soc. 1964,23, xlvi. Contributions to Books Mierobiology of digestion. With O. P. Cuthbertson. Chapter in the Biochemistry and Physiology 01 Nutrition, vol. 2, pp. 128-161. Academic Press, New York. 1953. Rumen dysfunction. Chapter contributed to Recent Advances in Veterinary Sciel1ce, vol. 2, pp. 213-260. Academic press, New York. 1955. The sheep. With W. Thomson. Chapter contributed to The UFA W Handbook on the Care and Management 01 Laboratory Animals. 2nd ed. Chapter 51, pp. 538-599. U.F.A.W., 7a Lamb's Conduit Passage, London, W.c.1. The alimentary tract of the ruminant. With D. Benzie. Monograph. Oliver & Boyd, Edinburgh. 1958. Metabolie disorders. With J. H. Bouckaert. Chapter 2 of Control 01 Diseases in Cattle and Sheep at Pasture, pp. 41-56. European Productivity Agency of O.E.E.C., Paris. 1958. The nutritional physiology of the adult ruminant. Chapter contributed to Animal Health Production and Pasture. Longmans, Green & Co., London. 1963. The digestion and absorption ofnitrogenous compounds in the ruminant. Chapter contributed to Mammalian Pro tein Metabolism, vol. 1, pp. 71-103. Academic Press, New York. 1964. Rumination ami the propulsion offood. Chapter contributed to Progress in Nutrition and Allied Sciences. Ed. D. P. Cuthbertson. Oliver & Boyd, Edinburgh. 1963. Veterinary studies. Chapter contributed to University Choice. Penguin Books Ud, Harmondsworth. June 1966. Ruminant digestion. Chapter contributed to Physiology 01 Domestic Animals, 8th ed. Dukes. Cornell University Press. Ch. 22. 1955. pp. 424-483. Absorption in the rumen. With A. Oobson. Chapter contributed to Handbook 01 Physiology 01 the Alimentary Canal (1968). Ameriean Physiologieal Society. Sect. 6, Vol. 5, Ch. 132, pp. 2761-2774.

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DIGESTlVE PHYSIOLOGY AND METABOLISM IN RUMINANTS

mieroflora of the alimentary tract of herbivora with special reference 10 ruminants. A. T. Phillipson. Nutr. Abst. Revs. 1947, 17, 12-18.

Physiological Mechanisms affecting the flow of digesta in Ruminants. With R. W. Ash. Chapter contributed to Physiology 0/ Digestion in the Ruminant. Ed. R. W. Dougherty. Butterworths, Washington. 1965. (Paper presented at 2nd International Symposium, Physiology of Digestion in the Ruminant, Ames, Iowa 1964).

Ruminant digestion. S. R. Elsden and A. T. Phillipson. Ann. Rev. Bioehem. 1948,17, 705-726. Absorption from the alimentary tract of sheep. A. T. Phillipson. Collected Papers, Rowett Research Institute 1955, 11, 31-37.

The nitrogenous compounds passing to the duodenum of sheep. With G. M. Ellinger. Chapter contributed to The Role 0/ the Gastrointestinal Traet in Protein Metabolism, pp. 137-142. Ed. H. N. Munro. 1964. (A symposium organised by the Council for International Organisations of Medical Sciences, Oxford: Blackwells) The metabolic diseases of ruminants. Chapter contributed to Nutrition o/Animals 0/ Agricultural Importanee, 17, part 11. Ed. D. F. Cuthbertson. Pergamon Press, Oxford. 1969. Absorption from the ruminant forestomach. With A. Dobson. Chapter contributed to Handbook Physiology, Section 6, Alimentary Canal. Ed. C. Code Amer. Physiol. Soe. 1969. Physiology of Digestion and Metabolism in the Ruminant. Proceedings of the Symposium on the subject held in Cambridge, August 1969. (A. T. Phillipson Chairman of Editorial Board.) Oriel Press, Newcastle-upon-Tyne. 1970. Ruminant digestion. Chapter 22 (pp. 424-484) of the 8th edition of Dukes' Physiology 0/ Domestie Animals. Cornell University Press, Ithaca and London. 1970. Ruminant digestion. Chapter 22 of the 9th edition of Dukes' Physiology 0/ Domestie Animals. Ed. M. J. Swenson. Comstock Publishing Assoc., Cornell University Press, Ithaca. 1977.

Miscellaneous Publications (Symposia, Lectures, Papers to Societies, etc.) Digestion in the ruminant. Ph.D. Thesis, Cambridge University. 1942. The pathogenesis of tympanites in cattle and the various gases involved. Veto Ree. 1942, 54, 430. Hypoglycaemia in the pregnant ewe (clinicaI communication). With M. W. S. Hitchcock. Veto Ree. 1946,58,504. The process of digestion in the ruminant. Br. Sei. News. 1948,1,8-10. Digestion in the ruminant. Nat. Advis. Serv., Q. Rev. No. 2, 55-62. 1948. Experimental ketosis in pregnant ewes. A. T. Phillipson. Symposium of toxaemias of pregnancy. Ciba Foundation Pub!. pp. 94-105. J. & P. Churchill Limited, London. 1950. The host in relation to alimentary organisms. A. T. Phillipson. Proe. R. Soe. B. 1952, 139, 196-201. (Part of a Symposium on Symbiosis involving micro-organisms.) Digestion of cellulose by the ruminant. A. T. Phillipson. Biochemical Society Symposia No. 11, 1953, 63-69.

Reviews Digestion in the ruminant. R. A. McAnally and A. T. Phillipson. Biol. Rev. 1944, 19,41. The physiology of digestion in the ruminant. A. T. Phillipson. Veto Ree. 1946, 58, 82. The fermentation of carbohydrate and the metaboIism ofthe derived fatty acids. Part 3 of a symposium entitled The role 0/ the

Digestion in the ruminant. A. T. Phillipson. X Vth International Veto Congress Proc., Pt. 1, Stockholm. Vo!. 1953, pp. 518-524. Resorptie door het Maagdarmstelsel van het Schaap. A. T. Phillipson. Vlaams Diergeneeskundig Tijdschrift. 1954, 23, 147-156.

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IN APPRECIATION OF ANDREW TINDAL PHILLIPSON De bewegingen yen de tractus digestions bij de herkanivers. A. T. Phillipson. Vlaams Diergeneeskundig Tijdschrift. 1954, 23, 113-122. (4) and (5): Two lectures delivered in the Veterinary School of the University of Ghent.

The nutritional physiology of the ruminant. A. T. Phillipson. Lecture at the Centenary of the Royal Veterinary and Agricultural College, Copenhagen. 1959. Printed in the College Yearbook 1959, pp. 7-16. The rumen in relation to the anima!. Proc. Nutr. Soc. 1959,18, 131-134.

Sodium transport and its role in ruminant digestion. A. T. Phillipson. (Paper delivered to the Animal Congress of the National Veterinary Medical Association.) Veto Rec. 1955,67, 1048-1051. Rumen dysfunction. A. T. Phillipson. Adv. Veto Sei. 1955,2, 212-261. Modern concepts of ruminant digestion and metabolism. A. T. Phillipson and D. P. Cuthbertson. (Paper delivered to the VIIth International Congress of Animal Husbandry.) Proc. 7th Int. Congress of Animal Husbandry, Madrid, 1956, pp. 7-93. Recent advances in ruminant digestion. A. T. Phillipson. Proc. 7th Int. Grassland Congress, New Zealand, 1956, 229-289. Ruminant digestion. A. T. Phillipson. Veto Rec. 1957,69, 11. Findings on pregnancy toxaemia in ewes. Agric. Rev. 1957, 3, 37-40. Bloat. Contribution to a symposium on grassland diseases organised by the European productivity Agency of the Organization for European Economic Cooperation. Project 204. 1958. Digestion in the ruminant. National Advisory Service QuarterI)' Review. 1958. No. 40, 1-11. Sheep. In Seientific Prineiples 0/ Feeding Farm Livestock. Proc. Conf. Brighton, 1958, pp. 105-119.. Farmer & Livestock Publications Ltd., London. 1959. Heritable characteristics of the red blood cells of sheep. With J. V. Evans. N.Z. Veto J. 1958,6, 12-14. Physiological effects of dis tension of the rumen and oesophagus in ruminants. With R. N. B. Kay. Proc. R. Soc. Med. 1959, 52, 374-376.

The nutrition of the ruminant. A. T. Phillipson. (Peter Wilson Bequest Lecture at the Royal (Dick) School of Veterinary Studies, July 1960). Vet Rec. 1960,72,613-616. The general physiology of the ruminant. A. T. Phillipson. In Digestive Physiology and Nutrition 0/ the Ruminant. Ed. D. Lewis. Butterworth, London. 1961. (Proceedings of the University of Nottingham 7th Easter School in Agricultural Science, 1960.) The nutrition of sheep. In Natl. Sheep Breeders Ass. Yearbook, pp. 34-38, 40. Radlett: C. R. Roberts. 1961. Nutrition as a factor in disease. A. T. Phillipson. Symposium on The Factors involved in the Study of Host/Agent Relationships. (Dedication of National Animal Diseases Laboratory, Ames, Iowa, U.S.A. December 12th and 13th, 1961. The flow of gastric contents to the duodenum of sheep. A. T. Phillipson and L. E. VowIes. 22nd Internat. Congr. Physio!. Sei. Leiden. Abstr. 397, 1962. Individuality and the effect of early summer grazing on ruminants. A. Dobson and A. T. Phillipson. In Animal Health and Production. Ed. C. S. Grunsell and A. r. Wright. Butterworths, London. 1962. (Proceedings of 13th Symposium of Colston Research Society, University of Bristol, 1961). Digestive system of the ruminant.

J. R. Agric. Soc. 1962,123,89-98.

The nitrogenous compounds passing to the duodenum of sheep. G. M. Ellinger and A. T. Phillipson. In The Role 0/ the Gastrointestinal Tract in Protein Metabolism. B1ackwell, Oxford. 1964, 137-142. (Proceedings of Symposium of The Council for International Organization of MedicaI Sciences, University of Glasgow, 1963.)

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DIGESTIVE PHYSIOLOGY AND METABOLISM IN RUMINANTS Comments on absorption in ruminants. A. T. Phillipson. Contribution to a symposium on Comparative Pharmacology in Washington. 1967. Fed. Proc. 26(4), 1006. Animal nutrition in the use and management programme. Contribution to a symposium on The Part the International Biological Programme will play in increasing World Jood supplies. Proc. Nutr. Soc. 26, 136, 1967. A. T. PhiIlipson. Fundamental research in animal science. A. T. PhiIlipson. Inaugural lecture given at the Portuguese Society of Veterinary Sciences, Nov. 1968. Rev. Port. Cienc. Veter. LXIV, No. 411. 1969.

The effects of diet and pentagastrin on the influx of urea into the rumen of sheep. C. J. F. Harrop and A. T. Phillipson. Proc. Nutr. Soc. 1971,30, 3A. Endogenous Josses of nutrients. A. T. Phillipson. Proc. Nutr. Soc. 1971,30,61-6. Intake, appetite and the fulfilment of alimentry requirements. A. T. Phillipson. Proceedings 2nd World Congress of Animal Feeding, Madrid. General Reports, 1972, pp. 703-707. The protection of dietary components from rumen fermentations. A. T. Phillipson. Proc. Nutr. Soc. 1972,31

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Acknowledgements The Scientific Committee and the local Organizing Committee of the Symposium would like to thank the Institut National de la Recherche Agronomique and the Ministere de I'Agriculture for their great assistance in preparing this conference. They would also like to thank the Vice-ChanceBor of the Universite de Clermont-Ferrand 1, the Dean of the Facultes de Medecine et de Pharmacie, and the Municipality of Clermont-Ferrand for the facilities they have aB put at the Organizing Committees' disposal. They wish to thank Le Centre National de la Recherche Scientifique, la Delegation Universitaire aux Relations Internationales, and the European Economic Community for their financial aid.

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Sponsors The following private firms and cooperatives have demonstrated their interest in the symposium by making financial contributions. The Scientific and Local Organizing Committees are very grateful to these sponsors. INTERNATIONAL SPONSORS BASF, FR Germany; Degussa, FR Germany; Upjohn International Inc., USA; Cyanamid International Corporation, Switzerland; Unilever, UK; Imperial Chemical Industries, UK; A. Coppens, Belgium; Bayer, FR Germany; British Petroleum, UK; J. Bibby Agriculture Ud., UK; Scottish Agricultural Industries Ud., UK. NATIONAL SPONSORS UTA; Eli Lilly, France; Hoffmann-La Roche et Cie.; Proligo; IBM France; Air France; Duquesne-Purina; Specia; Alfa-Laval Elevages; Univor; les Salins du Midi; Roussel-Uclaff; CERNA; Centrale Cooperative de Productions Animales (CCPA); Ucanor; Iffa-Merieux; Union des Cooperatives Agricoles d'Alimentation du Betail (UCAAB); Union Laitiere Normande (ULN); Roquette Freres; Avebene; Merck Sharp et Dohme, France; Tenstar Aquitaine; France Luzerne; GlEERNA; Laboratoire de Recherches et de Construction Electroniques (LRCE); Protector, France; Sanders; Guyomarc'h; Westfalia-Separator, France; Lorraine-Cottibar; SA l'Aurore; BEL; Lactina; Omnium Scientifique Industriel (OSI); Touzart-Matignon; Sopropeche. REGIONAL SPONSORS Banque Populaire de l'Auvergne et de la Correze; Produits Richemonts; Limagrain; Cooperative Cantalienne d'Elevage de l'Insemination Artificielle (CCEIA); Manufacture Fran~aise de Pneumatiques Michelin; Fressinet; Societe de Chimie Organique et Biologique, Commentry; Porcentre; Limoujoux Auvergne; Sucrerie de Bourdon; Grande Blanchisserie du Centre; Maillere. xxvi