0 and Digestive Physiology in Monogastric Farm Animals
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Nutrition ^ v; 0 anddigestiv e physiology inmonogastri c farmanimal s Reviews presented at the symposium on the occasion of the retirement of Dr Ir E.J. van Weerden, Wageningen, Netherlands, 26 May 1989 E.J. van Weerden & J. Huisman (Editors) Pudoc Wageningen 1989 !-"' i~* \ T>Y\ CIP data, Royal Netherlands Library, The Hague Nutrition anddigestiv e physiology in monogastric farm animals. Reviews presented at the symposium onth e occasiono f the retirement of Dr Ir E.J.va nWeerden ,Wageningen , Netherlands, 26 May 1989. / E.J. van Weerden &J. Huisman (Editors). - Wageningen: Pudoc. - x+IOI pp. ISBN 90-220-1011-2 UDC 636.2/.5.084:59l.l32 NUGI 835 Subject heading: animal nutrition © Centre for Agricultural Publishing & Documentation (Pudoc), Wageningen, Netherlands, 1989 All rightsreserved .Nothin gfro m thispublicatio n mayb ereproduced ,store d ina computerize d system or published in any form or in any manner, including electronic, mechanical, reprographic or photo graphic, without prior written permission from the publisher, Pudoc, P.O. Box 4, 6700 AA Wage ningen, Netherlands. The individualcontribution s inthi spublicatio n andan yliabilitie s arisingfro m them remainth e responsi bility of the authors. Insofar as photocopies from thispublicatio n arepermitte d byth e Copyright Act 1912,Articl e I6Ban d Royal Netherlands Decree of 20Jun e 1974(Staatsbla d 351) asamende d in Royal Netherlands Decree of 23Augus t 1985(Staatsbla d47 1) an db yCopyrigh t Act 1912,Articl e 17,th e legallydefine d copyright fee for anycopie sshoul db etransferre d to the StichtingReprorech t (P.O.Bo x 882, 1180 AW Amstel veen, Netherlands). For reproduction of parts of this publication in compilations such asanthologie s or readers (Copyright Act 1912, Article 16), permission must be obtained from the publisher. Printed in the Netherlands. BIBLIOTHEEK1 LANDBOUWUNIVERSITEIT ' ''.GENINGEN ^ÉÉf •FF1? V-."äjaft».ATt Si. - . -r i«.t J Dr Ir E.J. van Weerden PREFACE Dr Ir E.J. vanWeerde n together with Dr Ir P. van derWal , founded the institute now known as ILOB. Dr vanWeerde n started work there in 1960 andwa s responsible for nutritional and physiological research until 1984whe n ILOB became part of the TNO organization. Dr van der Wal then left ILOB.Upo n the departure of Dr van derWal ,D r van Weerden became also responsible for the general management of ILOB. TNO is an organization for applied scientific research and ILOB is an institute within TNO responsible for animal nutrition and physiology. It is partly subsidised by the government,mainl y through the Ministry of Economic Affairs, but the major part of thewor k is concerned with applied contract research. This is often confidential, implying inman y cases that the results can only be published with some delay; in some cases, no publications are allowed atall . Research at ILOB is concerned with various topics, such as: efficacy and safety of rawmaterials ,antibiotics ,probiotics ,ne w growth- promoting agents such as steroid hormones,/2-agonist san d somatropin; digestive physiology and nutrition ofmonogastri c farm animals, pharmacokinetics and biotransformation of additives and veterinary drugs. Dr.va nWeerde n had the ultimate responsibility for this research. Topics inwhic h hewa s especially involved were amino acid requirement in poultry, pigs and veal calves,digestiv e physiology in pigs and veal calves,efficac y and safety of single-cell protein, new growth-promoting agents,antibiotics , probiotics, replacement of skimmed milk powder by other protein sources. / Dr vanWeerde n is an excellent scientist, internationally recognized as a specialist in the fields mentioned who, in the period after 1984, also proved himself to be a good manager.Alway s sensitive to both sense and nonsense in research, he did much of the preliminary work on any new techniques or fields of research himself to seewhic h points may be critical. Hewa s invited to present his work atman y scientific congresses and had considerable influence onman y new developments in the fields of digestive physiology and nutrition of farm animals, advising various industries on the development of new products of application in the feed industry. A symposiumwa s organised inhonou r of Dr vanWeerde n to acknowledge all the effort he had put into the research carried out at ILOB for twenty-nine years and intomanagemen t for the last five years. Two specialists from abroad were invited to present papers on digestive physiology in pigs and veal calves.Tw o of his co-workers contributed papers on developments in amino acids and antinutritional factors.A paper on the effects of somatropin in pigs, resulting from the research that has been done for several years in cooperation with the Agricultural University, was also present. The symposiumwa s closed by Dr vanWeerde n with the presentation of his paper: Past and Future Developments of Protein Supply inMonogastri c FarmAnimals . In this book, the papers presented are published in an extended form as reviews. These contributions are just a small expression of the appreciation felt for Dr vanWeerde n by his colleagues and co-workers from ILOB. The organisers wish to thank Mr Haak,Mr s van den Berg-Volders and Mrs Haak-van den Brink for their excellent assistance in the organization. J. Huisman, J.B. Schutte, S. Bakker and G.J.M,va n Kempen (Organizing Committee) Wageningen, Netherlands,Ma y 1989. CONTENTS Research into the digestive physiology of pigs A.G. Low Antinutritional factors (ANFs) in the nutrition ofmonogastri c farm animals J. Huisman 17 Research into the digestive physiology of the milk-fed calf R. Toullec, P. Guilloteau 37 Effect of porcine somatotropin onnitroge n gain and energy metabolism in fattening pigs M.W.A. Verstegen,W . van der Hel 57 Practical application of (bio)synthetic amino acids in poultry and pig diets J.B. Schutte / Present and future developments in the protein/amino acid supply in monogastric farm animals E.J. vanWeerde n °9 RESEARCHINT OTH EDIGESTIV EPHYSIOLOG YO FPIG S A.G.LO W Pig Department,Agricultura l and Food Research Council Institute for Grasslandan dAnima lProduction ,Shinfield ,Reading ,Berkshir eRG 29AQ , U.K. Summary Historical development ofdigestiv e physiology is reviewed briefly. The importance of food intake,an d inparticula r its neuro-endocrine relationship,i sdiscussed . Gastric,biliar yan dpancreati csecretion s and the effects of age and diet are described. The function of the smallintestin ebot hi nterm so fsecretio nan dabsorptio nar ediscusse d with particular reference topractica lmeasuremen t ofprotei nqualit y and carbohydrate digestion and fermentation. The role of the large intestine in nutrient absorption is also discussed. Methods of measuring absorption using blood flow and sampling techniques are discussed. Therevie wend swit hsom estrategie sfo rfutur eresearch . Keywords / Pig,Digestion ,Secretion ,Absorption ,Stomach ,Smal lIntestine , LargeIntestine ,Protein ,Carbohydrate . 1. Introduction On the occasion of the retirement of E.J. Van Weerden after a distinguished career, it seems appropriate to review research in digestive physiology in thepig ,becaus e he hasbee n instrumental in several importantdevelopment s inth efield . Atth esam etim eh eha s encouraged others topursu e innovative studies and he has put TNO in theforefron to fresearc ho nthi ssubject . Historically, research into thedigestiv ephysiolog y ofpig swa si n progressa tRothamste d Research Station,Harpenden ,i nEnglan d inth e middleo fth enineteent hcentury ,whe nobservation swer emad eo nusin g anearl yfor mo fcannula . However,majo rdevelopment si nthi sfiel di n pigswer estil lnearl yon ehundre dyear saway . Inth elas tpar to fth e nineteenth century and theearl ypar t of this century Pavlov andhi s colleagues inRussi alai d thefoundation so f thesubjec to fdigestiv e physiology as we know it today. Their many experiments on the physiologyo f food consumption indogs ,an d its reflexbasis ,an dth e subsequentserie so fdigestiv ehydrolyses ,involve dth edevelopmen to f methodso f surgeryan d physiological principleswhic hwer e appliedt o pigs by Kvasnitskii and others from the 1930s onwards. Curiously enoughthes e studiesexcite d little interest inothe r countriesunti l the early 1960s when Horszczaruk and colleagues in Poland began to examinehindgu tfunctio ni npigs . Soonafterward sCunningham ,Frien d& Nicholson (1963)bega nt omak estudie si nCanad afollowe db yth eFrenc h groupi nJouy-en-Josa sle db yRéra t(Auffray ,Martine t& Rérat ,1967) , andth eEnglis hgrou pa tShinfiel d formedb yBraud edurin gth e1960s . Fromthe nonward sdevelopmen twa srapi di nal lo fthes ecountries ,an d during the 1970s, further active new groups were formed in the USA, Australia,Cuba ,Belgium ,th eNetherlands ,Denmark ,Sweden ,th eGerma n Democratic Republic and West Germany. Since 1979 there have been meetings on digestive physiology in the pig in Shinfield, England (1979), Jouy-en-Josas, France (1982), Copenhagen, Denmark (1985)an d Jab^onna,Polan d (1988). Thenex tmeetin gwill ,appropriately ,b ehel d during 1991 in Wageningen. Each meeting has been followed by a publication containing important reviews and papers on new research topics. The aim of this review is todiscus s selected topicswhic h have a particular interest for the author,an d tomak e some suggestions for futuredevelopment si nthi sfield . 2. Foodintak e Thecontro lo f food intakei npigs ,a si nman yothe ranimal san di n man, remains a topic of major interest and much uncertainty. The neuro-endocrinecontro lo ffoo dintak ei sbecomin gprogressivel ybette r understood,a s reviewsb yHoup t (1982)fo r thepig ,an d Forbes (1988) foranimal s ingenera lhav eshown . Butmuc h remainst ob eunderstoo d of what determines the pattern of food intake in relation to the physical, chemical and sensory properties of foods. While we know somethingo fth ewa yi nwhic hth ecentra lnervou ssyste mi sintegrate d for the processes of digestion, through the action of regulatory peptides,glucostati c andgastrointestina l controls,i ti snonetheles s the case that most studies have been made under physiological conditionswhic har edistan t fromthos ewhic h occurwhe nnorma lmeal s areeaten .