Dahlem Workshop Reports Life Sciences Research Report 25 Population Biology of Infectious Diseases The goal of this Dahlem Workshop is: to evaluate what the population biology of infectious agents implies for control of disease Life Sciences Research Reports Editor: Silke Bernhard Held and published on behalf of the Stifterverband fUr die Deutsche Wissenschaft Sponsored by: Senat der Stadt Berlin Stifterverband fUr die Deutsche Wissenschaft Population Biology of In fectious Diseases R. M. Anderson and R. M. May Editors Report of the Dahlem Workshop on Population Biology of Infectious Disease Agents Berlin 1982, March 14 -19 Rapporteurs: P.EM. Fine' M.P. Hassell, B.R.Levin· K.S.Warren Program Advisory Committee: R. M. Anderson and R. M. May, Chairpersons' J. Berger J. E Cohen' K. Dietz· E G. Knox· M.S. Pereira Springer-Verlag Berlin Heidelberg New York 1982 Copy Editors: M. Cervantes-Waldmann, K McWhirter Photographs: E.P.Thonke With 4 photos, 12 figures, and 14 tables ISBN-13: 978-3-642-68637-5 e-ISBN-13: 978-3-642-68635-1 001: 10.1007/978-3-642-68635-1 CIP-Kurztitelaufnahme der Deutschen Bibliothek: Population biology of infectious diseases: report of the Dahlem Workshop on Population Biology of Infectious Disease Agents, Berlin 1982, March 14-191 R. M. Anderson and R. M. May, ed Rapporteurs: P. E. M. Fine ... [Dahlem Konferenzen. Held and pub!, on behalf of the Stifterverb. fUr d. Dt. Wiss. Sponsored by: Senat d. Stadt Berlin; Stifterverb. fur d. Dt. Wiss.] - Berlin; Heidelberg; New York: Springer, 1982 (Life sciences research report; 25) (Dahlem workshop reports) NE: Anderson Roy M. [Hrsg.] Fine, Paul E. M. [Mitverf.]; Workshop on Population Biology of Infectious Disease Agents <1982, Berlin West>; Dahlem Konferenzen; GT This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law, wherre copies are made for other than private use, a fee is payable to "Verwertungsgesellschaft Wort", Munchen. © D. Bernhard, Dahlem Konferenzen, Berlin 1982 Softcover reprint of the hardcover 1s t edition 1982 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. 2131/3014 - 5 4 3 2 1 0 Table of Contents The Dahlem Konferenzen •.................••...•.•.....••.•. vii Introduction R.M. May ...•............•••...••...•.............••..••••• Impact of Infectious Diseases on Host Populations Group Report M.P. Hassell, Rapporteur R.C. Anderson, J.E. Cohen, B. CVjetanovic, A.P. Dobson, D.E. Gill, J.C. Holmes, R.M. May, T. McKeown, M.S. Pereira, D.A.J. Tyrrell •••..•............ 15 Impact of Infectious Disease Agents on the Population Growth and Geographical Distribution of Animals J .C. Holmes............................................... 37 The Impact of Infectious Disease on Human Demography Today M. S. Pereira.............................................. 53 Transmission: Patterns and Dynamics of Infectious Diseases Group Report K.S. Warren, Rapporteur R.M. Anderson, V. Capasso, A.D. Cliff, K. Dietz, F. Fenner, R.N. T-W-Fiennes, Z. Grossman, H. Knolle, P.G. Mann, L. Molineaux, G.A. Schad, D. Schenzle ••••.••.•• 67 Overall Population Patterns in the Transmission Cycle of Infectious Disease Agents K. Dietz.................................................. 87 Transmission Cycles and Broad Patterns of Observed Epidemiological Behavior in Human and Other Animal Populations F. Fenner ..•..•..•.......•......••.••.•..........•...••••• 103 vi Table of Contents Control of Infectious Diseases G1"OUp Repo1"t P.E.M. Fine, Rapporteur J.L. Aron, J. Berger, D.J. Bradley, H.J. Blirger, E.G. Knox, H.P.R. Seeliger, C.E.G. Smith, K.W. Ulm, P. Yekutiel ..................................... 121 Transmission Dynamics and Control of Infectious Disease Agents R.M. Anderson ............................................. 149 Practical Problems in the Control of Infectious Diseases C. E.G. Smith.............................................. 177 Use of Pathogenic Viruses As Agents for the Biological Control of Insect Pests L.A. Falcon .•...••...•••...•••.•.••.•....•..•.••..•.•••.•. 191 Evolution of Parasites and Hosts G1"OUp Repo1"t B.R. Levin, Rapporteur A.C. Allison, H.J. Bremermann, L.L. Cavalli-Sforza, B.C. Clarke, R. Frentzel-Beyme, W.O. Hamilton, S.A. Levin, R.M. May, H.R. Thieme •..••..•..•..•...•....•.• 213 Co-evolution Between Hosts and Infectious Disease Agents and its Effects on Virulence A.C. Allison .••.••••.•••..••.••••.•••..•••••••......••.... 245 Pathogens As Causes of Genetic Diversity in their Host Populations W.O. Hamilton •••••..•.•.••••••••.••.•.••••••••..••.•••••.. 269 Glossary ••.•.••.•.•.••.••••.••.•.•...••..•.....•..•..••••. 297 List of Participants...................................... 305 Subject Index ••.••••..••.••.•..•.•.••.•.•••.•.•••.••••..•. 309 Author Index .•••••••.•••..•.••••••••.••.•.•.•.•••••••••••. 315 The Dahlem Konferenzen DIRECTOR: Silke Bernhard, M.D. £:Q~I2?:,!!Q~.:. Dahlem Konferenzen was founded in 1974 and is supported by the Stifterverband fur die Deutsche Wissenschaft*, in cooperation with the Deutsche Forschungsgemeinschaft** and the Senat of the City of Berlin. Q12.z~~'!!~~:.. The task of Dahlem Konferenzen is: - to promote the interdisciplinary exchange of scientific in formation and ideas, - to stimulate international cooperation in research, and - to develop and test different models conducive to more effec- tive scientific meetings. AIM: Each Dahlem Workshop is designed to provide a survey of the present state of the art of the topic at hand as seen by the various disciplines concerned, to review new concepts and tech niques, and to recommend directions for future research. TOPICS: The workshop topics (in the Life Sciences and the field of Physicochemistry) should be: - of contemporary international interest, - timely, interdisciplinary in nature, and - problem-oriented. PROCEDURE: ---------- Dahlem Konferenzen approaches internationally recognized scien tists to suggest topics fulfilling these criteria and to pro pose members for a Program Advisory Committee, which is re sponsible for the workshop's scientific program. Once a year, the topic suggestions are submitted to a scientific board for approval. * The Donors Association for the Promotion of Sciences and Humanities **German Science Foundation viii The Dahlem Konferenzen g~~,!:!~!g~~,!§:.. The number of participants is limited to 48 for each workshop. They are selected exclusively by a Program Advisory Committee. Selection is based on international scientific reputation alone and is independent of national considerations, although a bal ance between Europeans and Americans is desirable. Exception is made for younger German scientists for whom 10% of the places are reserved. ,!~~_Q~~~~~-~Q~~§~Qg-~QQg~:.. A special workshop model has been developed by Dahlem Konferenzen, the Va.hlem Wolt/v.,hop MadeL The main work of the workshop is done in four small, interdisciplinary discussion groups, each with 12 members. Lectures are not given. Some participants are asked to write background papers providing a review of the field rather than a report on individual work. These are circulated to all participants 4 weeks before the meeting with the request that the papers be read and questions on them formulated be60lte the workshop, thus providing the basis for discussions. During the workshop, each group prepares a report reflecting the essential points of its discussions, including suggestions for future research needs. These reports are distributed to all participants at the end of the workshop and are discussed in plenum. g!!!!~!~~,!!Q~:.. The Dahlem Workshop Reports contain: - the Chairperson's introduction, - the Background Papers, and - the Group Reports. The Dahlem Workshop Reports are available in two series: 1) Life Sciences Research Reports (LS) and 2) Physical and Chemical Sciences Research Reports (PC). DAHLEM KONFERENZEN, Wallotstrasse 19, 0-1000 Berlin 33, Federal Republic of Germany Population Biology of Infectious Diseases, eds. R.M. Anderson and R.M. May, pp. 1-12. Dahlem Konferenzen 1982. Berl in, Heidelberg, New York: Springer-Verlag. Introduction R. M. May Biology Dept., Princeton University Princeton, NJ 08544, USA Old Peasant Many a man stands living here That your father, in the nick of time, Snatched from the fever's burning rage When he put limits to the plague. Faust My father was a good man lost in the dark. With what enthusiasm, what extravagance- Yet in his own odd way, in all sincerity- He brooded on Nature and her sacred soheres! He locked himself in the black kitchen With the other adepts; then with interminable Recipes, compounded incompatibles. Oh, he is happy who has any hope Of rising, ever, from this sea of error! What we don't know is just what we could use, And what we do know is no use to us. Goethe: Faust, Part One (Translation by R. Jarrell (5)) These lines are from the opening scenes of Goethe's great story. After receiving the veneration of the people as he moves among them at the fair, Faust confides to his companion, Wagner, that neither he nor his father had any real understanding of what 2 R.M. May they were doing when they fought against the plague; indeed, Faust says, he fears they killed more than they saved. Thus the stage is set for Faust's contract with the devil, as he sells his soul in exchange for knowledge. In the closing scenes of the drama, Faust's last gift to his people is to drain the marshes around the town -- a fully contemporary public health measure based on his dearly-bought understanding. The quest for clearer insight into the workings of disease in human and other animal populations runs as a theme throughout the history of biological science. The present time is one of substantial progress on many fronts. Advances in immunology and molecular biology are leading to a deeper understanding of the processes of infection within individual hosts; these and other advances are leading to new and more effective vac cines and drugs. Research on the evolutionary ecology and popu lation dynamics of intermediate vectors sheds light on some past failures in control programs and is helping in the design of more effective strategies of intervention. Mathematical epidemiology provides statistical tools for the reduction and analysis of data, along with an expanding literature on the factors governing the transmission and maintenance of endemic and epidemic infections. As can so easily happen in our age of increasing specialization, the cross-linkages between these various fields of endeavor are - with some notable exceptions -- not as strong as is desirable. Some of the mathematical literature has taken on a life of its own, free from data and full of elegant theorems in hopeful search of a disease. Conversely, some control programs have been based mainly on intuition, when a more analytic underpin ning could have been used. In particular, I think it is impor tant that our increasingly good understanding of the nature of immunity in individuals be matched with a clear appreciation of the processes of herd immunity. Even with cheap, safe, and effective vaccines or drugs, such understanding of immune pro cesses at the level of the population as a whole is necessary
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