Contemporary Topics in Immunobiology: Self/Non-self Discrimination PDF
Preview Contemporary Topics in Immunobiology: Self/Non-self Discrimination
CONTEMPORARY TOPICS IN IMMUNOBIOLOGY 9 VOLUME CONTEMPORARY TOPICS IN IMMUNOBIOLOGY General Editor: M. G. Hanna, Jr. Frederick Cancer Research Center Frederick, Maryland Editorial Board: Max D. Cooper University of Alabama Birmingham, Alabama John J. Marchalonis Frederick Cancer Research Center Frederick, Maryland G. J. V. Nossal The Walter & Eliza Hall Institute ofM edical Research Victoria, Australia Victor Nussenzweig New York University School of Medicine New York, New York George W. Santos Johns Hopkins University Baltimore, Maryland Ralph Snyderman Duke University Medical Center Durham, North Carolina Osias Stutman Sloan-Kettering Institute for Cancer Research New York, New York Noel L. Warner University of New Mexico Albuquerque, New Mexico William O. Weigle Scripps Clinic and Research Foundation La Jolla, California A Continuation Order Plan is available for this series_ A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher. CONTEMPORARY TOPICS IN IMMUNOBIOLOGY 9 VOLUME Self / Non -self Discrimination EDITED BY 1. JOHN MARCHALONIS Frederick Cancer Research Center Frederick, Maryland and NICHOLAS COHEN University of Rochester Rochester, New York PLENUM PRESS • NEW YORK AND LONDON The Library of Congress cataloged the first volume in this series as follows: Contemporary topics in immunobiology. v. 1- 1972- New York, Plenum Press. v. illus. 24cm. annual. 1. Immunology-Periodicals. QR180.C632 574.2'9'05 79-179761 ISSN 0093-4054 Library of Congress Catalog Card 79-179761 ISBN 978-1-4615-9133-7 ISBN 978-1-4615-9131-3 (eBook) DOl 10.1007/978-1-4615-9131-3 © 1980 Plenum Press, New York Softcover reprint of the hardcover 1s t edition 1980 A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N.Y. 10011 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher F. Macfarlane Burnet first used the phrase "self/non-self discrimination" in 1940. His concepts have provided a chal lenge to two generations of immunologists. It is with great pleasure that we dedicate this volume to Sir Mac on the oc casion of his 80th birthday. J. J. Marchalonis N. Cohen M. G. Halma, Jr. Contributors Thomas C. Cheng Institute for Pathobiology Center for Health Sciences Lehigh University Bethlehem, Pennsylvania 18015 Michael J. Chorney Institute for Pathobiology Center for Health Sciences Lehigh University Bethlehem, Pennsylvania 18015 Adrienne E. Clarke School of Botany University ofM elbourne Parkville, Victoria 3052, Australia Nicholas Cohen Department of Microbiology Division of Immunology University of Rochester School of Medicine and Dentistry Rochester, New York 14642 Jack D. Cowan Department of Biophysics and Theoretical Biology and the College University of Chicago Chicago, minois 60637 Michael Edidin Department of Biology The Johns Hopkins University Baltimore, Maryland 21218 B. F. Edwards Department of Biology Reed College Portland, Oregon 97202 Charles A. Janeway, Jr. Department of Pathology Immunology Division Yale University School ofM edicine New Haven, Connecticut 06510 R. Bruce Knox School of Botany University of Melbourne Parkville, Victoria 3052, Australia vii viii Contributors Heinz Kohler La Rabida-University of Chicago Institute Chicago, illinois 60649 and Departments ofP athology and Biochemistry University of Chicago Chicago, illinois 60637 Stephen P. Lerman Department ofP athology New York University School ofM edicine New York, New York 10016 John J. Marchalonis Cancer Biology Program NCI Frederick Cancer Research Center Frederick, Maryland 21701 Daniel Meruelo Irvington House Institute Department of Pathology New York University Medical Center New York, New York 10016 Michael A. Palladino Department of Pathology New York University School ofM edicine New York, New York 10016 Donald A. Rowley La Rabida-University of Chicago Institute Chicago, Illinois 60649 and Departments ofP athology and Pediatrics University of Chicago Chicago, Illinois 60637 L. N. Ruben Department of Biology Reed College Portland, Oregon 97202 G. Jeanette Thorbecke Department of Pathology New York University School ofM edicine New York, New York 10016 Gregory W. Warr Cancer Biology Program NCI Frederick Cancer Research Center Frederick, Maryland 21701 Preface The problem that virtually all cells have in discriminating between "self" and "non-self" molecules and cells has been considered at great length in immuno biology. However, cells that clearly are incapable of carrying out mammalian type immune functions can exhibit exquisite specificity in their capacity to discriminate among syngeneic, allogeneic, and xenogeneic cells. In this volume of Contemporary Topics in Immunobiology we have chosen to consider the general problem of self/non-self discrimination as it is manifest in recognition reactions of plants and invertebrates and in the evolutionary development of the immune response of vertebrates. A broad, many-faceted approach is taken toward fundamental issues in immunobiology in order to develop innovative concepts of receptor function as well as to delineate traditional views. The capacity of plants to discriminate between self and non-self is addressed in Chapter 1 by R. B. Knox and Adrienne E. Clarke. These authors provide examples of cell-cell recognition in plants that parallel those occurring in in vertebrates and vertebrates. In general, tolerance (acceptance) of grafts is re stricted to plants within closely related genera. Recognition is mediated by callus cells, which proliferate at wound surfaces in higher plants, and there is a correlation between cell and tissue type and antigenic markers detectable with the use of mammalian antibodies. Certain flowering plants exhibit precise discrimination in fertilization, when pollen must be from the same species, but fertilization occurs only if the pollen is genetically non-self. Self-incompatibility provides an excellent model for study of the genetic and molecular basis of recognition in plants because a specific recognition, or S-gene, system has been described and an antigenic glycoprotein has been implicated as the S-gene product. The authors discuss cell-cell recognition in plants in the context of concepts derived in cell biology and cellular immunology. M. J. Chorney and T. C. Cheng in Chapter 2 consider cellular recognition reactions in invertebrates. They focus upon three questions: (1) Do invertebrates show specificity in the recognition of antigen? (2) What is the molecular basis for this interaction? (3) Does a phylogenetic lineage of immunoreactive com plexity exist, which eventually leads to the complex vertebrate immune response? ix x Preface They discuss phagocytic-cell reactions and humoral factors such as agglutinins and consider the possible roles of lectins and putative precursors of immunoglob ulins and major-histocompatibility-complex (MHC) products in the elaboration of specific cell recognition by invertebrate cells. L. N. Ruben and B. F. Edwards (Chapter 3) and G. Jeanette Thorbecke, M. A. Palladino, and S. P. Lerman (Chapter 4) address the evidence for the exis tence and function of cell-cell cooperation in specific immune responses of nonmammalian vertebrates. Ruben and Edwards provide an extensive review of data pertinent to the existence of primary and secondary lymphoid organs and lymphoid heterogeneity in lower vertebrates (cyclostomes, fishes, amphibians, and reptiles). They propose that collaborative cellular interactions in humoral immunity may have been a fundamental feature of vertebrate evolution, and that the appearance of such interactions did not await the evolutionary emer gence of the bone marrow or bursa of Fabricius as hematopoietic sources. They present evidence supporting the provocative concept that complexity of T-cell function increases with evolution within each vertebrate class and suggest that the primitive lymphocytes in evolution might resemble mammalian B cells rather than T cells. Thorbecke et al. give a detailed, scholarly review and present new evidence relating to lymphoid-cell cooperation in immune responses of the chicken, a species that has been used effectively in recent studies of T and B cells. These authors provide clear evidence that T cells have the capacity to exhibit a fine degree of molecular recognition specificity, and they further define the complexity of T-B and T-macrophage interaction in this avian species. The importance of the MHC in many immunological phenomena has led to research into the evolutionary history of this gene complex. N. Cohen (Chapter 5) analyzes recent immunological and immunogenetic studies with cold-blooded vertebrates and suggests that MHC phylogeny may be a saga of convergent evolution of Class-II-region genes (i.e., I-region homologous). Functional markers characteristic of this region in mice indicate the presence of the MHC in advanced fishes and anuran amphibians and its absence in primitive fishes, primitive amphibians, and some, if not all, reptiles. Salamanders have been the most extensively studied group of vertebrates that apparently lack the complete homologue of the MHC. Although a mixed-lymphocyte-reaction (MLR) locus has been provisionally identified in salamanders, it is not a major-histocompatibility (class-I) locus and it is not functionally linked to a predominant H locus. More over, the salamander system described by Cohen is of considerable interest to those attempting to ascertain theoretically the function of the MHC, because the putative MLR locus is minimally polymorphic, possessing only two codomi nant alleles. G. W. Warr (Chapter 6) confronts the evidence for the existence of immuno globulins on the surface of lymphocytes of vertebrate species ranging from Preface xi teleost fishes to mammals. He marshals clear evidence supporting the presence of immunoglobulin polypeptide determinants on T- and B-like lymphocytes of representatives of all classes of vertebrates studied, and proposes that immuno globulin, or a structurally related molecule, serves as the receptor for primary binding of antigen by all lymphocytes in all vertebrates. Monomeric IgM, or a closely related immunoglobulin, appears to be universally present as an antigen receptor on lymphocytes of the B-cell series. In addition, a second major class of B-Iymphocyte surface immunoglobulin (IgD) has been described in primates and some rodents. T cells bear immunoglobulinlike receptors whose heavy chains are likely to be of a new and unique class, and the association of light chains with these molecules is a controversial issue. The preceding chapters document the evolutionary generality (within the vertebrates) of cell-cell cooperation, the importance of the MHC, and the uni versality of immunoglobulin in antigen recognition. C. A. Janeway, Jr., (Chapter 7) concentrates on well-studied rodent models and addresses the complexity of T-cell populations and the function of distinct T-cell subsets in positive and negative (suppressive) T-T, T-B, and T-macrophage interactions. He emphasizes the importance of the MHC for specific recognition of antigen by T cells and offers a provocative hypothesis that accounts for the requirement for products of immunoglobulin genes (VH) and MHC genes in self/non-self discrimination by T cells. He hypothesizes that T cells carry at least two kinds of specific receptors, one for MHC antigens and one for non-MHC antigens. Both of these receptors are predicted to bear Vwencoded and distinct idiotypic determinants. D. A. Rowley, H. Kohler, and J. A. Cowan (Chapter 8) approach the prob lem of immunoregulation at its highest level of complexity. They develop the concept of an "immunologic network" in which the intensity and quality of immune responses arise from a complex regulatory system consisting of mole cules and cells specifically connected by the complementarity of receptors for antigen and receptors for receptors. The network is modulated by multiple positive and negative feedback loops. D. Meruelo and M. Edidin (Chapter 9) offer a novel approach to the bio logical function of the MHC by clarifying and unifying all of the processes and interactions that appear to be regulated or modified by the MHC. Besides the involvement of the MHC in the genetic control of immune responses, the MHC exerts some genetic influence over a variety of traits not currently considered to have an immunological basis. These include the following: levels of plasma testosterone, weights of steroid-sensitive organs, mating behavior, develop mental effects, e.g., cleft palate, levels of cAMP, binding of hormones to cell membranes, and cell to cell adhesion. Meruelo and Edidin build upon these observations to develop a broadly based concept of MHC function alternative to those proposed on the basis of immunological studies (for example, see Chapter 7). They suggest that the products of the MHC, especially serologically
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