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Lectu re Notes in
,B iomathematics
Managing Editor: S. Levin
32
Systems Theory
in Immunology
Proceedings of the Working Conference
Held in Rome, May 1978
Edited by
Carlo Bruni, Gino Doria, Giorgio Koch, and Roberto Strom
Springer-Verlag
Berlin Heidelberg New York 1979
Editorial Board
w. Bossert· H. J. Bremermarm . J. D. Cowan· W. Hirsch· S. Karlin
J. B. Keller· M. Kimura· S. Levin (Managing Editor) . R. C. Lewontin
R. May· G. F. Oster· A. S.Perelson . L. A. Segel
Editors
Carlo Bruni Gino Doria
Istituto di Automatica Chen-Euratom Immunogenetics Group
University of Rome Laboratory of Radiopathology
Via Eudossiana 18 C.S.N. Casaccia (Rome), Italy
00184 Rome, Italy
Giorgio Koch Roberto Strom
Istituto Matematico Istituto di Chimica-Biologica
"G. Castelnuovo" Facolta di Medicina e Chirurgia
University of Rome University of Rome
Rome, Italy 00185 Rome, Italy
AMS Subject Classifications (197~: 92-06
ISBN-13: 978-3-540-09728-0 e-ISBN-13: 978-3-642-93130-7
001: 10.1007/978-3-642-93130-7
This work is subject to copyright. All rights are reserved, whether the whole
or part of the material is concerned, specifically those of translation, re
printing, re-use of illustrations, broadcasting, reproduction by photocopying
machine or similar means, and storage in data banks. Under § 54 of the
German Copyright Law where copies are made for other than private use,
a fee is payable to the publisher, the amount of the fee to be determined by
agreement with the publisher.
© by Springer-Verlag Berlin Heidelperg 1979
PREFACE
This volume collects the contributions presented at the "Working Conference on
System Theory in Immunology", held in Rome, May 1978.
The aim of the Conference was to bring together immunologists on one side and
experts in system theory and applied mathematics on the other, in order to identify
problems of common interest and to establish a network of joint effort toward their
solution.
The methodologies of system theory for processing experimental data and for
describing dynamical phenomena could indeed contribute significantly to the under
standing of basic immunological facts. Conversely, the complexity of experimental
results and of interpretative models should stimulate mathematicians to formulate
new problems and to design appropriate procedures of analysis.
The multitude of scientific publications in theoretical biology, appeared in
recent years, confirms this trend and calls for extensive interaction between mathe-
matics and immunology.
The material of this volume is divided into five sections, along the scheme of
the Conference program.
The articles in the first section deal with experimental and theoretical aspe
cts of the mechanisms of antigenic stimulation of immunocompetent cells. The second
section discusses the mechanisms of cell interactions in the regulation of antibody
production. Section three contains contributions on the problems of the evaluation
of antibody affinity and of the maturation of the immune response. The last two
sections are devoted to immunological modeling. In particular, the papers in section
four consider the po tentiality of mathematical modeling to dissect out the varia
bles involved in immune functions and/or to reach a comprehensive view of the whole
system. Biological and mathematical aspects of network theory are throughly examined.
in section five.
The editors wish to express their gratitude to the University of Rome and to
the Italian National Research Council (C.N.R.) for having sponsored the Conference,
and to the Accademia Nazionale dei Lincei under whose auspices the Conference was
held. They are also grateful to Springer-Verlag, for publishing the Proceedings of
the Conference in the Lecture Notes in Biomathematics.
Rome, September 1979
Carlo Bruni
Gino Doria
Giorgio Koch
ltoberto Strom
CONTRIBUTORS
Riehard Asofsky Laboratory of Microbial Immunity National Institutes of
Health. Bethesda, Maryland.
Mario Barra Istituto Matematico G.Calstelnuovo - University of Rome.
Rome, Italy.
Erie B. BeU Department of Experimental Pathology - Manchester Univer
sity Medical School. Manchester, England.
George I. BeU Los Alamos Scientific Laboratory-University of California.
Los Alamos, New Mexico.
Dietmar G.Braun Institute for Immunology and the Biocenter - University
of Basel . Basel, Switzerland.
Carlo Bruni Istituto di Automatica - University of Rome.
Rome, Italy.
Franeo Celada Department of Immunology - University of Genoa.
Genoa, Italy.
Valeriano Comineioli Laboratorio di Analisi Numerica - Consiglio Nazionale
delle Ricerche. Pavia, Italy.
Gabriella Del Grosso Isituto M.atematico G.Castelnuovo - University of Rome.
Rome Italy.
Charles De Lisi Laboratory of Theoretical Biology - National Cancer Insti-
tude. Bethesda, Maryland.
Gino Doria CNEN - Euratom Immunogenetics Group-Laboratory of Radio-
pathology of C.S.N. Casaccia. Rome, Italy.
Alberto Gandolfi Centro di Studio dei Sistemi di Controllo e Cal colo
Automatici - Consiglio Nazionale delle Ricerche.
Rome, Italy.
VI
Anna Gerardi Istituto Matematico G.Castelnuovo - University of Rome.
Rome, Italy.
Alfredo Germani Centro di Studio dei Sistemi di Controllo e Calcolo
Automatici - Consiglio Nazionale delle Ricerche.
Rome, Italy.
Maria Adelaide Giovenco Centro di Studio dei Sistemi di Controllo e Calcolo
Automatici - Consiglio Nazionale delle Ricerche.
Rome, Italy.
Zvi Grossman Laboratory of Theoretical Biology-National Cancer Institute
Bethesda, Maryland.
Raymond Hamers Institute voor MoleculaireBiologie - Vrije Universiteit.
Brussels. St.Genesius-Rode, Belgium.
Jacques R.Hiernaux Laboratory of Theoretical Biology - National Cancer
Institute. Bethesda, Maryland.
Geoffrey W. Hoffmann Basel Institute for Immunology.
Basel, Switzerland.
Sarah Howie Basel Institute for Immunology.
Basel, Switzerland.
Chin-Shung Hsu Department of Electrical Engineering -
Cleveland State University. Cleveland, Ohio.
Giorgio Koch Istituto Matematico G.Castelnuovo - University of Rome.
Rome, Italy.
Ignace Lasters Instituut voor MoleculaireBiologie-
Vrije Universiteit Brussel. St.Genesius-Rode, Belgium.
Oberdan Leo Department de Biologie Moleculaire -
Universite Libre de Bruxelles. St.Genesius-Rode, Belgium.
VII
Federico Marchetti Istituto Matematico G.Castelnuovo -
University of Rome. Rome Italy.
Rona"ld R.Moh"ler Department of Electrical and Computer Engineering -
Oregon State University. Corvallis, Oregon.
Goran MaUer Department of Immunobiology-
Karolinska Institute, Wallenberglaboratory.
Stockholm, Sweden.
MuUer Moser Department de Biologie Moleculaire -
Universite Libre de Bruxelles. St.Genesius-Rode, Belgium.
Luigi NespoU Laboratorio di Analisi Numerica -
Consiglio Nazionale delle Ricerche. Pavia, Italy.
Arduino Ora tore Istituto di Chimica Biologica -
Istituto Universitario di Medicina e Chirurgia.
L'Aquila, Italy.
Pierfrancesco Periti Laboratorio di Analisi Numerica -
Consiglio Nazionale delle Ricerche. Pavia, Italy.
Christian P"lasschaert Department de Biologie Moleculaire -
Universite Libre de Bruxelles. St.Genesius-Rode, Belgium.
Peter H.Richter Department of Chemistry -Massachusetts Institute of
Technology. Cambridge, Massachusetts.
Wo"lfgang Scha"lch Basel Institute for Immunology.
Basel, Switzerland.
Anne"liese Schimp"l Institut fur Virologie und Immunbiologie.
University of Wurzburg. Wurzburg, W.Germany.
Giuseppe Serazzi Laboratorio di Analisi Numerica -
Consiglio Nazionale delle Ricerche. Pavia, Italy.
VIII
Roberto Strom Istituto di Chimica Biologica-
University of Rome. Rome, Italy.
Hans P.Tony Institut fur Virologie und Immunbiologie-
University of Wurzburg. Wurzburg, W.Germany.
Jacques J.Urbain Laboratorie de Physiologie Animale-
Universite Libre de Bruxelles. St.Genesius-Rode, Belgium.
PhiZippe Van Ham Logical and Numerical Systems Department-
Bruxelles University. Bruxelles, Belgium.
Eberhard Wecker Institut fur Virologie und Immunbiologie-
University of Wurzburg. Wurzburg, W. Germany.
Keith J.Wright Max-Planck-Institut fur Biologie.
Tubingen, W. Germany.
Christian WUiZmart Department de Biologie Moleculaire-
Universite Libre de Bruxelles. St.Genesius-Rode, Belgium.
CONTENTS
I. Antigenic Stimulation
Mechanism of B Lymphocyte Activation. 2
G. MoHer
Clonal Dominance: Presentation of Antigenic Subsites is 13
the Determining Factor.
D.G. Braun. W. SahaLah. J.K. Wright
Hierarchic Immunogenicity of Protein Determinants. 28
F. CeLada
Control of B Lymphocyte Activation through Reversible 37
Binding of Multivalent Antigen: a Simple Model.
A. GandoLfi. M.A. Giovenao. R. Strom
II. Cell Interactions 53
Some Cellular Interactions Involved in the Primary Immune 54
Response.
S. Howie
Some Models for the Interaction between Cells of the 66
Immune Systems.
G.I. BeH
Stimulation of Memory Cells and Abrogation of Suppression 75
with a Mobile Antigen - laden Cell.
E.B. BeLL
Involvement of the Fc Receptor in Immune Regulation: 87
a Hypothesis.
A. SahimpL. H.P. Tony. E. Weaker
III. Evaluation and Evolution of Antibody Production 103
Problems in the Evaluation of Antibody Affinity Distri 104
bution during the Immune Response.
R. Strom. C. Bruni. A. Germani. G.Koah. A. Oratore
