Biology of the Mouse Histocompatibility-2 Complex

Transcription

1 Biology of the Mouse Histocompatibility-2 Complex

2 Peter A. Gorer (Courtesy of Mrs. P. A. Gorer) George D. Snell (Courtesy of Dr. G. D. Snell)

3 Biology of the Mouse Histocompatibility-2 Complex Principles of Immunogenetics Applied to a Single System Jan Klein Springer-Verlag New York Heidelberg Berlin 1975

4 Jan Klein Department of Microbiology The University of Texas Southwestern Medical School 5323 Harry Hines Boulevard Dallas, Texas Library of Congress Cataloging in Publication Data Klein, Jan, Biology of the mouse histocompatibility-2 complex. Bibliography: p H-2 locus. 1. Title. [DNLM: 1. Chromosomes. 2. Histocompatibility. 3. Immunogenetics. QH600 K64bl. QRI84.3.K54 599' ISBN-13: e-isbn-13: : / All rights reserved. No part of this book may be translated or reproduced in any form without written permission from Springer-Verlag by Springer-Verlag New York Inc. Softcover reprint of the hardcover I st edition 1975

5 With love to my mother, and with apologies to Dagmar, Norman, and Daniel.

6 "Ach, wenn man doch wissend werden konnte!" rief Knecht. "Wenn es doch eine Lehre gabe, etwas, woran man glauben kann! Alles widerspricht einander, alles lauft aneinander vorbei, nirgends ist GewiBheit. Alles labt sich so deuten und labt sich auch wieder umgekehrt deuten... Gibt es denn keine Wahrheit? Gibt es keine echte und guitige Lehre?" Hermann Hesse, Das Glasperlenspiel

7 Preface Somewhere I heard a story of a bridge and a painter. The bridge was enormous and was made all of metal, and the painter's job was to keep it from rusting. He would start at one end and slowly proceed, day by day, month by month, toward the other end, painting the bridge. But no sooner would he finish with the painting than the bridge would begin to rust again. The rust, too, would start at one end and slowly proceed toward the other end, systematically destroying the painter's endeavor. And so the painter would return to where he had started, and begin painting again, slowly proceeding toward the other end of the bridge, always just one step ahead of the rust. And if the story is true, the painter might still be painting that bridge-a modern Sisyphus! During the writing of this book, the story of the painter and his bridge kept coming to mind. The field the book covers has been developing so rapidly that, like the painter, I too had to return to where I had started and fight the rust of obsolescence. But unlike the painter, I had a deadline to meet, which constituted a point of no return. And so, sending off this manuscript, I have no choice but to watch the fruits of my endeavor be overtaken by the rust. In recent years, the H-2 system has become not only the prototype of similar systems in other species, but also one of the focusing points of modern biology. It forms a bridge between immunology and genetics, between biochemistry and immunology, between virology and genetics, and it carries the potential of broad biological implications. I have tried to cover all these different facets of the system and to present as complete a picture as was in my power. The breadth of this book's scope forced me to plunge into areas in which I am not an expert and thus increase the risk of having errors slip into the text. Although I tried to minimize such risk through consultations with many of my colleagues, I probably have not avoided it completely. For any errors remaining, I sincerely apologize. Anticipating that the book might be read by geneticists who know little about immunology, immunologists who know little about genetics, and others who are neither immunologically nor genetically oriented, I have placed at the beginning of each section an introductory chapter summarizing the basic knowledge necessary for understanding the particular section. Because of the broad area covered and because of the limited space available, I could not document fully by references many important observations mentioned in this monograph. This is particularly true for the introductory chapters. As a rule I have deliberately avoided references to work done in species other than the mouse. This too was motivated only by space limitations. ix

8 x Preface I am indebted to many individuals who, in different ways, assisted me during the work on this monograph: to Dr. Donald C. Shreffler, with whom I spent my Lehrjahre; to Drs. James Forman, Dietrich G6tze, Margaret C. Green, Frank Lilly, Eva Lotzova, Hugh O. McDevitt, George D. Snell, and Ellen S. Vitetta who read portions of the book and made invaluable comments and suggestions; to all those quoted in the text who provided me with unpublished data and important documentary material; to Messrs. Jong M. Park and Suresh Savariryan, whose outstanding technical expertise allowed me to continue my research while working on this book, and to Springer-Verlag for having the courage to undertake this venture. But above all, I thank Ms. Pamela Erbe Wiener, who so patiently struggled with my distortions of her native tongue. Jan Klein

9 Contents Preface ix Chapter One: Chapter Two: History Part One: Introduction The mouse and its forms 3 16 Part Two: Peripheral Regions of the H-2 Complex Section One: Serology Chapter Three: Chapter Four: Chapter Five: Introduction to serological analysis Methods of serological H-2 typing Current status of H-2 serology Section Two: Histocompatibility Chapter Six: Chapter Seven: Chapter Eight: Introduction to histocompatibility Methods of histocompatibility testing Histocompatibility system of the mouse Section Three: Genetics Chapter Nine: Chapter Ten: Chapter Eleven: Chapter Twelve: Chapter Thirteen: Principles of genetic analysis Recombination and mutation in the H-2 complex Genetics of H-2 linked loci Population genetics of the t and H-2 systems The H-2 complex and somatic cell genetics Section Four: Biochemistry Chapter Fourteen: Cellular and subcellular distribution of H-2 antigens Chapter Fifteen: Biochemical aspects of H-2 antigens xi

10 xii Contents Part Three: Central Regions of the H-2 Complex Section Five: I-region associated traits Chapter Sixteen: Genetic control of virus susceptibility 389 Chapter Seventeen: Genetic control of immune response 411 Chapter Eighteen: Genetic control of cell recognition and interaction 449 Section Six: S and X region-associated traits Chapter Nineteen: The S region and miscellaneous H-2-associated loci 493 Part Four: Conclusion Chapter Twenty: Function and evolution of the H-2 complex 52 I Appendix: Chronology of major events in the history of the H-2 system 537 References 541 Author Index 603 Subject Index 611