CELLULAR ASPECTS OF IMMUNITY Ciba Foundation Symposia General Volumes : Mammalian Germ Cells - - - - - Preservation and Transdantation of Normal Tissues - - - - - - LeukaemiaResearch - - - - Chemistry and Biology of Pteridines - Porphyrin Biosynthesis and Metabolism Histamine - - - - - - Extrasensory Perception - - - Bone Structure and Metabolism - - Paper Electrophoresis - - - - Ionizing Radiations and Cell Metabolism The Nature of Viruses - - - - Chemistry and Biology of Purines - - Drug Resistance in Micro-organisms - Chemistry and Biology of Mucopolysaccharides The Cerebrospinal Fluid- - - - - Neurological Basis of Behaviour - - - Amino Acids and Peptides with Antimetabolic Activity - - - - - - - Medical Biology and Etruscan Origins - - Biosynthesis of Terpenes and Sterols - - Carcinogenesis: Mechanisms of Action - - Regulation of Cell Metabolism - - - Biochemistry of Human Genetics - - - Significant Trends in Medical Research - - Congenital Malformations - - - - A leajlel giving details of the Ciba Foundation Colloquia on Endocrinology, Colloquia on Ageing and Study Group volumes is available from the Publishers CIBA FOUNDATION SYMPOSIUM ON CELLULAR ASPECTS OF IMMUNITY Editors for the Ciba Foundation G. E. W. WOLSTENHOLME, O.B.E., M.A., M.B., M.R.C.P. and MAEVE O’CONNOR, B.A. With 11 7 Illustrations LITTLE, BROWN AND COMPANY BOSTON Library of Congress Catalog ('nrd Number: 00-11 885 ALL RIGHTSR 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 Publishers. THE Ciba 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 Foundation offers accommodation to scientists from all over the world at its home in Portland Place. Foremost in its acti- vities 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 in- formal discussions between research workers of different dis- ciplines and different nationalities and each year invites an outstanding authority to deliver a special lecture. An exchange programme between French and British postgraduates is con- ducted 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 grad- uates 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 this ancient Swiss town. V THE CIBA FOUNDATION for the Promofion of International Co-operation in Medical and Chemical Research PORTLAPNLDA CEL,O NDONW, .l. 41 Trustees THER IGHTH ONL. ORDA DRIAN,O .M., F.R.S. THER IGHTH ON.L ORDB EVERIDGEK,.C .H., F.R.A. SIR RUSSELLB RAINB, T. THEH ON.S IR GEORGEL LOYD-JACOB SIR RAYMONNDE EDHAMQ, .C., F.S.A. Executive Council SIR RUSSELBL RAINB, T., Chairman THEH ONS. IR GEORGEL LOYD-JACOB PROFESSAO. RH ADDOWF.,R .S. PROFESSODRR . DR. h.c. R. MEIER SIR ARTHURV ERE HARVEY, PROFESSFO. RG . YOUNGF,. R.S. C.B.E., M.P. Director, and Secretary to the Executive Council DR. G. E. W. WOLSTENHOLMEO, .B.E. Assistant Secretary MISS N. BLAND Scientific Assistant Editorial Assistant MISS CECILIAM . O'CONNOR,B .Sc. Miss MAEVEO 'CONNOR,B .A. Conference Assistant MRS. NANCYG . SPUFFORD vi PREFACE INi ts 10th Anniversary year the Ciba Foundation for the first time arranged three conferences away from its own premises in London. The second of these took place in the Abbey of Royaumont, now the property of the Cercle Culture1 de Royaumont, which lies in peaceful wooded country near Chantilly, 22 miles north of Paris. This volume is a record of the scientific proceedings of that meeting. Unfortunately no report can be given of the social events or do justice to the hospitality all participants received from the French members present, many of whom also worked hard in helping with the preparation and running of the symposium. The original suggestion for a conference on a subject con- nected with delayed hypersensitivity reactions came from Professor Kourilsky, after consultation with Professor Bug- nard, and it was with their invaluable advice and assistance, and that of Professors Oberling, Grabar and Medawar, that the Director of the Foundation was able to decide on the final title and programme of the meeting. Professor Kourilsky also agreed to act as Chairman, a task which he performed with much energy and discretion. At a later stage, when the book was being prepared for publication, Professor Medawar, Professor Miescher and Dr. Humphrey gave reassuring advice for which the editors would like to thank them. vii CONTENTS PAGE Chairman's opening remarks R. KOURILSKY. 1 Short Communication : Nomenclature of immunologically competent cells . by ASTRIDF AGRAEUS 3 An in vitro study of some mechanisms of antigen uptake by cells . by R. ROBINEAUaXnd JANINPrEN m 5 Discussion : DIXON, HALPERNH, UMPHREYL, IXIERBID~G, MIESCIIERR, OBINILWXSM, ITHV, OISIN . . 40 Degradation of a protein antigen by intracellular enzymes by C. LAPRESLaEn d T. WNBB . . 44 Discussion : FAGRAEUGSRA, BAR, HALPERN,H ARRISH, UM- PHREY, KUNKEL, LAP. RESLE,L EDERBICRGM, I.:DAWA.R , MILES, ROBINEAUX .57 Microcinematographic contributions to the study of plasma cells J. P. TIIIERY . . 39 by Ultrastructure of immunologically competent cells by W. BERNHARD and N. GRANBOULAN . . !,a nisLcEusDsiEoRn B: BICERRSNEGLII,IA GRMLA),N FNA, GSRIMIEOCNSS, ENG,O TOIDII, kRGYR. %B.' t\n, IhRRIS., 117 Identification of immunologically competent cells by M. SIMONSEN . . 122 Discussion : BURNETG, ORERI,I ARRISL,E DERBEILMGI,X MWAR, MITCIIISONS, IMONSEN. . 131 Theories of immunological tolerance by P. B. MEDAWAR . . 134 Discussion :B vitNm, DIXONG, OOD,G ORERG, RABARH,A RRIS, HUMPHREYL, EDERBERGM, E. DAW AR, MIESCIIER,M ILES. , MONOD,S MITHW, AKSMAN 149 viii CONTENTS ix PAGE Panel discussion on the clonal selection theory of antibody formation . . BURNETL, ED'CRB ERG 157 70ith DIXONG, OOD, (:RAB.lR, H~RRIJSO,I IANOVSKhTfE~D, AWAR, MILIIS, MONOII, SIMONSENS, rEnzL, VoIsrN Lymph node cell transfer in relation to antibody formation . . ly T. N. HARRIaSn d SUSANNAH ARRIS 172 Formation of antibodies in vitro by P. GRABARa nd P. CORVAZIER. . 198 Discussion : DIXONS, TERZI,F, AGRAEUS . . 206 A study of serum proteins related to immunity and their cellular origins by P. BURTIN . . 213 Discussion : BURTIN,G OOD, KUNKELM, I~.:SCITERM, ITCHISON, SF.I,IGM.4NN, SMITH . . 224 The cellular basis for the immunological memory . . by D. W. DRESSERan d N. A. MITCIIISON 227 Discussion : BURNET,D IXON, FAGRAEUGSO,R ER, GRABAR, HALPERNH, ARRIS,L EDERBEKG, MEDAWAR, MITCIIISON, MONOD,S IMONSEN, ShlITII . . 238 Some biological and immunological properties of transfer factor . by H. SIIERWOODL AWRENCE * ?43 Discussion : GOOD, JOHANOVLASWKRJE, NCEL,E DERBERG, . . MILES, SMITHW, AKSMAN 272 A comparative histopathological study of delayed hyper- sensitive reactions . . by B. €1. WAKSVAN 2 y Discussion : GOOD, Gnmnn, JOIUNOVSK~K.,O URILSKY, LEDERBERG, MEDAWAR, MILES, OBERLING, SIMONSEN, WAKSMAN . . 322 Interactions between sessile and humoral antibodies in homograft reactions . . by P. A. GORER 330 Discussion :B ERNHARGDO, RER,G RABARH, UMPHRELYE,D ER- . . BERG, MEDAWAR, MIESCHEH, OBERLING, WAKSMAN 343 X CONTENTS PAGE Response to active immunization of human infants during the neonatal period . . by T. SMITII R. 348, Discussion : FAGRAECSG, OOD,G ORER,K UNKEI,L, EDERBERG, MEDAWARS, ELIGMANNS,M ITH,S TERZL . . 369 Modifications of capillary permeability in immunological reactions mediated through cells by G. A. VOISINa nd FRANCITNOEU LLET . . 376 . Discussion : MILES, VOISIN,W AKSMAN 405 The patterns and mechanism of fixation of anaphylactic antibody on living tissues in vitro by B. N. HALPERN,P . LIACOPOULOSR, . BIN.~GHaIn d . R. LIAcoPouLos-BIiIor 409- Discussion :G RABARH, ALPERNH, UMPIIREYM, IESCHERM, ILES, . SMITII 425 Multiple “autoantibodies” to cell constituents in systemic . lupus erythematosus 4d by H. G.K UNKELH, . R. H&.MANan d H. R. G. DEICIIICR Discussion : Go~DK, UNKEI,, MIESCIIICRM, ONOD,R OBINEAUX, . . . SELIGMANN, STERZI,, WAKSMAN 438 The in vitro action of antigen-antibody complexes on thrombocytes and erythrocytes by P. MIESCHER,N . S. COOPERa nd D. HVREZ . . 4 6 Discussion : HCMPIIREYK,O URILSKMY,I ESCHER,H OBINEACX 459 Group discussion in summary BURNET,w ith DIXON, FAGRAEUS, GOOD, GHABAR, SUSANNAH ARRISH, UMPHREYK,U NREL,L EDERBERG, MEDAWARM, ILES,M ONOD,S IMONSENW, AKSMANSM, ITH, . . STERZL 461 . . Chairman’s closing remarks 475