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PART TWO THEORETICAL IMMUNOLOGY PART TWO THEORETICAL IMMUNOLOGY THE PROCEEDINGS OF THE THEORETICAL IMMUNOLOGY WORKSHOP, HELD JUNE, 1987 IN SANTA FE, NEW MEXICO Alan S. Perelson, Editor Theoretical Division Los Alamos National Laboratory m Volume SANTA FE INSTITUTE STUDIES IN THE SCIENCES OF COMPLEXITY Advanced Book Program CRC Press Taylor & Francis Group Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business Director of Publications, Santa Fe Institute: Ronda K. Butler-Villa First published 1988 by Westview Press Published 2018 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 CRC Press is an imprint of the Taylor & Francis Group, an informa business Copyright 1988 by Taylor & Francis Group LLC No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reason-able efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copy- right holders if permission to publish in this form has not been obtained. If any copy- right material has not been acknowledged please write and let us know so we may rec- tify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permis- sion from the publishers. For permission to photocopy or use material electronically from this work, please access www. copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organiza-tion that provides licenses and regis- tration for a variety of users. For organizations that have been granted a photocopy li- cense by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trade- marks, and are used only for identification and explanation without intent to infringe. Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Library of Congress Cataloging-in-Publication Data Theoretical Immunology Workshop (1987 : Santa Fe, N.M.) Theoretical immunology. (Santa Fe Institute studies in the sciences of complexity ; v. 2-3) Includes index. 1. Immunology--Mathematical models--Congresses. 2. Immunology--Philosophy--Congresses. 3. Immunologic diseases-- Mathematical models--Congresses. I. Perelson, Alan S., 1947- II. Title. 111. Series. [DNLM: 1. Allergy and Immunology--congresses. 2. Models, Theoretical—congresses. QW 504 T3955t 1987] QR182.2.M36T485 1987(cid:9) 616.079(cid:9) 88-6169 ISBN 0-201-15682-2 (pt. 1) ISBN 0-201-15683-0 (pbk.: pt. 1) ISBN 0-201-15687-3 (pt. 2) ISBN 0-201-15688-1 (pbk.: pt. 2) This volume was typeset using TEXtures on a Macintosh computer. ISBN 13: 978-0-201-15688-1 (pbk) About the Santa Fe Institute The Santa Fe Institute (SFI) is a multidisciplinary graduate research and teaching institution formed to nuture research on complex systems and their simpler elements. A private, independent institution, SFI was founded in 1984. Its primary concern is to focus the tools of traditional scientific disciplines and emerging new computer resources on the problems and opportunities that are involved in the multidisciplinary study of complex systems -- those fundamental processes that shape almost every aspect of human life. Understanding complex systems is critical to realizing the full potential of science, and may be expected to yield enormous intellectual and practical benefits. All titles from the Santa Fe Institute Studies in the Sciences of Complexity series will carry this imprint which is based on a Mimbres pottery design (circa A.D. 950-1150), drawn by Betsy Jones. Santa Fe Institute Series List Lecture Notes Volumes in the Santa Fe Institute Studies in the Sciences of Complexity Volume I: John Hertz, Anders Krogh, Richard G. Palmer, editors: Introduction to the Theory of Neural Networks Volume II: Gerard Weisbuch: Complex Systems Dynamics Volume III: Wilfred Stein and Francisco J. Varela, editors: Thinking About Biology Volume IV: Joshua M. Epstein: Nonlinear Dynamics, Mathematical Biology, and Social Science Volume V: H.F. Nijhout, Lynn Nadel, and Daniel L. Stein, editors: Pattern Formation in the Physical and Biological Sciences Proceedings Volumes in the Santa Fe Institute Studies in the Sciences of Complexity Volume I: David Pines, editor: Emerging Synthesis in Science Volume II: Alan S. Perelson, editor: Theoretical Immunology, Part One Volume III: Alan S. Perelson, editor: Theoretical Immunology, Part Two Volume IV: Gary D. Doolen, senior editor: Lattice Gas Methods for Partial Differential Equations Volume V: Philip W. Anderson, Kenneth J. Arrow, and David Pines, editors: The Economy as an Evolving Complex System Volume VI: Christopher G. Langton, editor: Artificial Life Volume VII: George I. Bell and Thomas G. Marr, editors: Computers and DNA Volume VIII: Wojciech H. Zurek, editor: Complexity, Entropy, and the Physics of Information Volume IX: Alan S. Perelson and Stuart A. Kauffman, editors: Molecular Evolution on Rugged Landscapes: Proteins, RNA, and the Immune System Volume X: Christopher G. Langton, Charles Taylor, J. Doync Farmer and Steen Rasmussen, editors: Artificial Life II Volume XI: John A. Hawkins and Murray Gell-Mann, editors: The Evolution of Human Languages Volume XII: Martin Casdagli and Stephen Eubank, editors: Nonlinear Modeling and Forecasting Volume XIII: Jay E. Mittenthal and Arthur B. Baskin, editors: The Principles of Organizations in Organisms Volume XIV: Daniel Friedman and John Rust, editors: The Double Auction Market: Institutions, Theories, and Evidence Volume XV: Andreas S. Weigend and Neil A. Gershenfeld, editors: Time Series Prediction: Forecasting the Future and Understanding the Past Volume XVI: George J. Gummerman and Murray Gell-Mann, editors: Understanding Complexity in the Prehistoric Southwest Volume XVII: Christopher G. Langton, editor: Artificial Life III Volume XVIII: Gregory Kramer, editor: Auditory Display: Sonification, Audification, and Auditory Interfaces Volume XIX: George A. Cowan, David Pines, and David Meltzer, editors: Complexity: Metaphors, Models, and Reality Volume XX: David Wolpert, editor: The Mathematics of Generalization Volume XXI: P.E. Cladis and P. Palffy-Muhoray, editors: Spatio-Temporal Patterns in Nonequilibrium Complex Systems Volume XXII: Harold J. Morowitz and Jerome L. Singer, editors: The Mind, the Brain, and Complex Adaptive Systems Volume XXIII: Bela Julesz and Ilona Kovacs, editors: Maturational Windows and Adult Cortical Plasticity Volume XXIV: Joseph A. Tainter and Bonnie Bagley Tainter, editors: Evolving Complexity and Environmental Risk in the Prehistoric Southwest Volume XXV: John B. Rundle, Donald L. Turcottc, and William Klein, editors: Reduction and Predictability of Natural Disasters Volume XXVI: Richard K. Belew and Melanie Mitchell, editors: Adaptive Individuals in Evolving Populations Volume XXVII: W Brian Arthur, Steven N. Durlauf, and David A Lane, editors: The Economy as an Evolving Complex System II Volume XXVIII: Gerald Myers, editor: Viral Regulatory Structures and Their Degeneracy Santa Fe Institute Publications Committee January, 1988 L. M. Simmons, Jr., Chair Vice President, Academic Affairs, Santa Fe Institute Dr. Robert McCormick Adams Secretary, Smithsonian Institution Professor Philip W. Anderson Department of Physics, Princeton University Dr. George I. Bell Leader, Theoretical Division, Los Alamos National Laboratory Dr. George A. Cowan President, Santa Fe Institute and Senior Fellow, Los Alamos National Laboratory Professor Murray Gell-Mann Division of Physics & Astronomy, California Institute of Technology Professor John H. Holland Division of Computer Science & Engineering, University of Michigan Dr. Bela Julesz Head, Visual Perception Research, AT&T Bell Laboratories Professor Stuart Kauffman School of Medicine, University of Pennsylvania Professor David Pines Department of Physics, University of Illinois

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