THE EXOCRINE PANCREAS Normal and Abnormal Functions Ciba Foundation Symposia General Volumes MammalianGermCells - - - - - 30s. Preservation and Transplantation of Normal Tissues - - - - - - - - 25s. LeukaemiaResearch - - - - - - 30s. Chemistry and Biology of Pteridines - - - 4s. Porphyrin Biospthesis and Metabolism - - 30s. H i s u e - - - - - - - - 50s. Extrasensory Perception - - - - 27s. 6d. Bone Structure and Metabolism - - - - 459. Ionizing Radiations and Cell Metabolism - - 45s. TheNatureofViruses - - - - - 49. Chemistry and Biology of Purines - - - 48s. Drug Resistance in Micro-organisms - - - 50s. TheCerebrospinalFluid - - - - - 50s. Amino Acids and Peptides with Antimetabolic Activity - - - - - - - - 459. Medical Biology and Etruscan Origins - - 45s. Biosynthesis of Terpenes and Sterols - - - 45s. Carcinogenesis: Mechanisms of Action - - 48s. Biochemistry of Human Genetics - - - 50s. Significant Trends in Medical Research - - 50s. CongenitalMalformations - - - - - 459. Haemopoiesis: Cell Production and its Regula- tion- - - - - - - - - 60s. AdrenergicMechanisms - - - - - 70s. Quinones in Electron Transport - - - - 60s. TheNatureofSleep - - - - - - 50s. Somatic Stability in the Newly Born - - - 50s. Renal Biopsy - - - - - - - 60s. EnzymesandDrug Action- - - - - 65s. Tumour Viruses of Murine Origin - - - 60s. A leajet giving details of the Ciba Foundation Colloquia on Endocrinology, Colloquia on Ageing and Study Group volumes is availablefrom the Publishers CIBA FOUNDATION SYMPOSIUM ON THE EXOCRINE PANCREAS Normal and Abnormal Functions Editors for the Ciba Foundation v. s. A. DE MUCK, M.SC., D.I.C., A.R.C.S. and MARGARET P. CAMERON, M.A. With 91 Illustrations J. and A. CHURCHILL LTD. 104 Gloucester Place, London, W.I 1962 ALLR IGHTSR ESERVED This book is protected under the Berne Convention. It may not be reproduced by any means, in whole or in part, without permission. Application with regard to reproduction should be addressed to the Pu blishers. Printed in Great Britain THEC iba Foundation, a unique international institution, owes its inception to the generosity of CIBA Limited, Basle. However, being established under British trust law, it enjoys complete independence in practice and policy. Under the guidance of its distinguished Trustees, the Founda- tion offers accommodation to scientists from all over the world at its home in Portland Place. Foremost in its activities is the organization of small conferences, the proceedings of which are published in book form in the manner of the present volume. The Foundation convenes many other informal discussions between research workers of different disciplines and different nationalities and each year invites an outstanding authority to deliver a special lecture. An exchange programme between French and British postgraduates is conducted and a library service is available. Furthermore, the Ciba Foundation attempts in every other way possible to aid scientists, whether they be Nobel Laureates or young graduates making their first original contribution to research. The purpose of the Ciba Foundation, which is to promote international co-operation in medical and chemical research, is symbolized in the armorial bearings by five interlaced rings representing the continents, a black sacrificial cock (emblem of Aesculapius) holding a medical caduceus, and three regular hexagons for chemistry. Its domicile in London is indicated by the red sword of St. Paul and the British lion; the wyvern and the crozier, symbols associated with Basle, refer to the sponsoring firm located in &IS ancient Swiss town. V THE CIBA FOUNDATION for the Promotion of International Cwperation in Medical and Chemical Research 41 PORTLANPDL ACBL, ONDONW, .l. Trustees THER IGHT HON.L ORDA DRIAN,O .M., F.R.S. THER IGHT HON. LORDB EVERIDGEK, .C.B., F.B.A. THER IGHT HONL. ORDB RAIN, U.D., D.C.L. THEH ON.S IR GEORGLEL OYD-JACOB SIR RAYMONDN EEDHAMQ,. C., F.S.A. Executive Council THER IGHTH ON.LO RD BEAIN,L L.D., Sm ARTHURV B~BH ARVECY.B,.E . D.C.L.,C hairman M.P. DR. F. GROSS THEH ON.S IR GEORGLEL OYD-JACOB PROPESSOAR. HADDOWF,. R.S. PROFESSOF.R G. YOUNGF,.R .S. Director, and Secretary to the Executive Council DR. G. E. W. WOLSTBNHOOL.MB.EE. , Deputy Director MR. A. v. s. DE REUCK, M.SC. Assistant Secretary Mrss N. BLAND Scientific Assistant and Librarian Editorial Assistant Mxss M. P. CAMERONM, .A. Mrss MAEVEO 'CONNORB, .A. Conference Rcsistant MRS. NANCYG . SPUPFORD PREFACE THISS ymposium arose out of a suggestion made by Dr. H. D. Janowitz, who pointed out that the Ciba Foundation had not hitherto devoted a meeting to any aspect of the alimentary tract. The scope and structure of the Symposium owe much to Dr. Janowitz’s generous advice and assistance with the organization of the meeting. All those connected with the Symposium would also wish to pay tribute to Professor Sheila Sherlock, who was persuaded to take the Chair, and whose skilful guidance contributed so much to the discussions recorded here. The meeting brought together a catholic range of specialists, including investigators who had been working on the morphology of the exocrine pancreas, thosc concerned with the processes of electrolyte and enzyme secretion by the gland, and also some concerned with the metabolic aspects of exocrine pancreatic disease. The membership was kept small, as at all Ciba Foundation meetings, in the interests of free and intimate discussion, but it is hoped that the publication of these proceedings wrll enable a much wider interested public to share to some extent in a most rewarding occasion, and that many may be stimulated by the general discussion at the end covering topics for future research- zymogen granules; the nervous control of pancreatic secretions ; pancreatic regeneration; standardization of tests ; the genetics of pancreatic disease; and pancreatic biopsy. vii CONTENTS Ultrastructure and Histochemistry PAGE The h es tructure of the exocrine pancreas cells by F. S. SJOSTRAND . . . . . . . . , I Discussion: BORGSTROMEK, HOLMG, ROSSMAHNA, RPERL, . E. HOKIN, HUNTJ,A NOWITZ, PALADES, HERLOCSKJ,O STFLWD. . I9 Structure, chemistry and function of the pancreatic exocrine cell by G. E. PALADPE. SIEKEVITZa nd L. G. CARO . . . . 23 Discussion: BORGSTROGMR, OSSMALN. ,E . HOKINM, . R. HOKINH, OWAT, PALADE, SJOSTRANn . . . . . . . . 49 Histochemistry of the exocrine pancreas byV. BECKER. . . . . . . . . . 56 Discussion: BECKERD, REILINGE, KHOLM, GROSSMANH,A RPERH,U NT, JANOWITLZA, GERLOPFA, LADEV, ~GHELYI . . . . . 6 4 Nature of Pancreatic Secretions Considerations of the occurrence, structure and function of the proteolytic enzymes of the pancreas by H. NEURATH . . . . . . . . . 67 Discussion: BORGSTROMD,E SNUELLGE,R OSSMALNA, GERLONF,E URATH, PALADE,TUNBRIDGE. . . . . . . . 86 Influence of the composition of the diet on the enzyme content of rat pancreas by P. DESNUELLJE. ,P . REBOUDa nd A. BENA BDELJLI.L . . g o Discussion: BORGSTROMBU, RNETT, DESNUELLDER, GILINGGR, OSSMAN, . L. E. HOKIN,H UNTJA,N OWITZ, LAGERLONFE, URATHP,A LADEV, m 107 The pancreatic secretion of fluid and electrolytes by H. D. JANOWITZ and D. A. DREILING . . . . 115 Discussion: BERNDB, JANOWITLZA, GERL~P . . . . I33 * Short Communication Relations between the volume and output of bicarbonate in pancreatic secretion by J. J. BERNIEaRn d A. LAMBLING . . . . . . I38 Discussion: BERNIERC, ARD, DREILING, HARPER, HUNT,J ANOWITZ, LAGBBLUP,NEURATE,V~G.HBL.~I . . I45 * * viii CONTENTS ix of Physiological Control Pancreatic Secretion - PAGE The gastrointestinal hormones secretin and cholecystokinin by J. E. JORPES and V. MUTT . . . . . . . 150 Discdon: CARDD, ESNUELLDER,E ILINGG, ROSSMAHNO, WATJA, NOWKTZ, JORPESM, UTT, SAIUES, SIE3RLOCK . . . . . . . 164 The distribution and physiological properties of pancreozymin by A. A. HARPERE., L. BLAIRan d T. SCRATCHERD . . . 168 Discussion: DREILING,G ROSSMAHNA, RPERL, . E. HOKIN,M . R. HOKIN, HUNTJ, ANOWITZLA, GERLOSFH, ERLOCK, VBGHJZYX . . . . I82 The synthesis and secretion of digestive enzymes by pancreas tissue in vitro by L. E. Horn and M ~ ERL. H orn. . . . . . 186 Discussion: DAWSONG, ROSSMAHNA, RPERL, . E. HOKIN,M . R. HOKIN, JANOWITPZA, LADES, JOSTRANVDI,N K . . . . , 204 Nervous and hormonal regulation of pancreatic secretion by M. I. GROSSMA.N . . . . . . . . 208 Discussion: DREILINGG, ROSSMANHA, RPER,M . R. HOKIN, HOWAT, LAGERLOSFH, ERLOCK . . . . . . . . . 220 Assessment of Pancreatic Function The measurement of pancreatic secretory function by D. A. DREILINaGn d H. D. JANOWITZ . . . . 225 Discussion: BURNETTD, REILINGG, ROSS,H OWAT,L AGERL~SFA, RLES, SHERLOCK . . . . . . . . . . 2 5 3 Normal and abnormal digestive function by G. LUNDHan d B. BORGSTR~M. . . . . . 259 Discussion: BORGSTRODMA, WSOND,R EILINGG,R OSSMAHNO, WATH, UNT, JANOWITLZA, GERLOSFA,R LES,S HERLOCVK~, GHEL.Y I . . . a74 Abnormal Pancreas: Genetic and Metabolic Aspects Further observations on the hereditary form of pancreatitis by J. B. GROSSJ., A. ULRICHan d F. T. MAHER . . . . 278 Discussion: BURNETDT,R EILINGG,R OSSG, ROSSMALN. E, . HOKINH, OWAT, JANOWITZ, MUTT,S HERLOCTKU, NBRIDVGBEG, HELYVI, INK . . 305 Diagnostic value of the sweat test in cystic fibrosis ~~C.L.J.VINK . . . . . . . . . 310 Diswssion: L. E. Horn, HOWATH, UNT,J ANOWKTMZU,T T, PALADE, SHERLOCK,V~GHELYIV, INR . . . . . . . . 325 X CONTENTS PAGE Protein metabolism and pancreatic function by P. V. V~GHELYanId T. T. KEM$NY . . . * . 329 Discussion: BORGSTR~DMR, EILPIGGR, OSSMANH,A IWm, L. E. Horn, . . LAGERL~NPE, URATSH,m ,S HE~OCISCJ,~ STRANVD~,G HELYI 349 General Discussion Zymogengranules . . . . . . . . . 353 Nervous control of pancreatic secretion . . . . . . 357 Pancreatic regeneration . . . . 358 Standardization of enzyme methods . . . 3 59 Standardization of function tests . . 362 Standardization of sweat tests. . 3 66 Genetics of pancreatic disease . . . . . . 36 7 Pancreatic biopsy . . . . . . . 370 BECKERB,E BNIERB, ORGSTR~BMUR, NETTC,A RDD,E SNUELDER, SILING, GROSSG, ROSSMAWN, ELR. E., H orn, M. R. Horn, HOWATH, UNT, JANOWITZ, JORPESL, AGERLOPM, m, NEURATPHAL, ADE, SAWS, SHERLOCISCJ,~ STRANVD~, GHELYVII,N K. Author index . . . . . . . 375 Subjectindex . . . . . . . 377
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