2001Basic SciencesBiological Sciences (Evolution, Behavior, Ecology, Environment)
John Maynard Smith photo

John Maynard Smith

  • U. K. / 1920-2004
  • Evolutional Biologist
  • Professor Emeritus, University of Sussex

Outstanding Contribution to Evolutionary Biology by Proposing the Idea of the Evolutionarily Stable Strategy (ESS)

Professor Maynard Smith has made a groundbreaking contribution to the establishment of a unified understanding of fundamental issues in evolutionary biology, including social activities of organisms and the essence of the evolution of sexual reproduction, by proposing the idea of the evolutionarily stable strategy (ESS). In doing so, he has not only immensely contributed to the development of biological sciences,but he has also had major impact on other disciplines including economics and politics.


Brief Biography

Born in London, England
B.A. in Engineering, University of Cambridge
Aircraft Engineer
B.Sc. in Zoology, University College London (UCL), University of London
Lecturer in Zoology, UCL
Professor of Biology, University of Sussex
First Dean, School of Biological Science, University of Sussex
Professor Emeritus, University of Sussex

Selected Awards and Honors

Fellow, The Royal Society
Darwin Medal, The Royal Society
Frink Medal, Zoological Society, London
Balzan Prize, Italy
The Linnean Medal, The Linnean Society, London
The ASAB Medal, The Association for the Study of Animal Behaviour,
U. K.
Foreign Members
The American Academy of Arts and Sciences
Honorary Doctors
University of Kent, University of Oxford, University of Chicago, University of Sussex, University of Edinburgh

Major Works

  • Evolution and the Theory of Games, Cambridge University Press, 1983

  • Evolutionary Genetics, Oxford University Press, 1989

  • The Major Transitions in Evolution, W. H. Freeman, 1995

  • Group selection and kin selection, Nature, 200, 1964

  • Evolution in sexual and asexual populations, The American Naturalist, 102, 1968

  • The logic of animal conflict. Nature, 246, 1973

  • Parental investment: a prospective analysis, Animal Behaviour, 25, 1977


Outstanding Contribution to Evolutionary Biology by Proposing the Idea of the Evolutionarily Stable Strategy (ESS)

Professor John Maynard Smith has made a groundbreaking contribution to the establishment of a unified understanding of fundamental issues in evolutionary biology, including the social behavior of organisms and the problems of sex and sex ratio, by proposing the idea of the Evolutionarily Stable Strategy (ESS).

According to the basic concept of modern biology, the behaviors of individual organisms and their efforts to reproduce are not directed towards the maintenance of their species, but are instead quite selfish and competitive, and are actually attempts to maximize their individual Darwinian fitness. i.e. to produce as many offspring with their own genes as possible. Altruistic and competitive behaviors appear to contradict this concept, but such interactions between related individuals have been satisfactorily explained by the concept of kin selection proposed by Professor W. D. Hamilton (a 1993 Kyoto Prize Laureate).

However, altruistic or cooperative behavior also often occurs between non-kin individuals; for example, when adult male animals fight for food or territory, they seldom fight to the death. Why? This question also very much pertains to human morality and peace, but it had hitherto eluded theoretical explanation.

Based on an assumption that animals are playing games for existence with each other, Professor Maynard Smith introduced game theory, and clearly demonstrated the reason why animals do not kill their competitors when they fight. He used the analogy of a Dove strategy versus a Hawk strategy: the Dove retreats at once if the opponent escalates the contest, whereas the Hawk always escalates the contest and continues until injured or until the opponent retreats. If all individuals are using a Dove strategy, a Dove strategy is more likely to increase individual fitness. However, individuals adopting the Dove strategy are always defeated by invading Hawk strategists, and so the population becomes unstable. Professor Maynard Smith proposed that there should be a strategy, termed an Evolutionarily Stable Strategy (ESS), which is more likely to succeed and persist in a population. Such a strategy shown by the majority of individuals in a population can effectively exclude invasion by any other strategy. He subsequently applied game theory to a wide variety of examples, to demonstrate how certain behaviors can be an ESS. For example, in the case of a fight, one ESS is a conditional strategy, which he termed a bourgeois strategy, saying `Play Hawk if owner of territory, and play Dove if intruder.’

Based on the idea of ESS, Professor Maynard Smith has used a mathematical approach to successfully and elegantly explain the essential aspects of a large number of unsolved questions in evolutionary biology, behavioral ecology and population genetics. As such, the theory of ESS is a key to a more unified understanding of the diverse field of biology. Furthermore, in addition to dramatic advances in the field of evolutionary biology, the ESS idea has also significantly influenced such diverse fields as economics, business sciences, and politics.

For these reasons, the Inamori Foundation is pleased to bestow upon Professor Maynard Smith the 2001 Kyoto Prize in Basic Sciences.


Abstract of the Lecture

The Marriage of Mathematics and Biology

I am being honoured for my contributions to evolutionary biology, and in particular for the introduction of Evolutionary Game Theory. This has required the marriage of two unrelated skills: knowledge of natural history, and skill at making mathematical models of the world. My love of nature has been with me since childhood, and continues to be an important part of my life, from bird-watching to gardening. My mathematical abilities were encouraged by excellent teaching at school, and a facility for applying mathematics to real problems was developed during six years working in aircraft design, and then by studying under J. B. S. Haldane. Evolutionary Game Theory was first developed to analyse the evolution of ritualized fighting in animals, but has since been applied to topics as diverse as plant growth and the evolution of viruses. My greatest happiness in science has come when some animal is found doing something odd, that was predicted by the theory.

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Evolution and Game

13:00-17:40, Monday, November 12, 2001
Kyoto International Conference Hall
Coordinator & Moderator:
Yoh Iwasa, Member of the Kyoto Prize Screening Committee; Professor, Department of Biology, Faculty of Sciences, Kyushu University


Opening Yoh Iwasa
Address Toyomi Inamori
Managing Director, The Inamori Foundation
Address Toshitaka Hidaka, Chairman, the Ktyoto Prize Comiittee in Basic Sciences; Director General, Research Institute for Humanity and Nature
Introduction of Laureate Yoh Iwasa
Laureate Lecture John Maynard Smith Laureate in Basic Sciences
"The Evolution of Animal Signals"
Lecture Tetsuo Kuwamura
Member of the Kyoto Prize Screening Committee; Professor, College of Liberal Arts, Chukyo University
"Game Theory for Animal Behavior"
Question - and - Answer
Lecture Norio Yamamura
Professor, Director of Center for Ecological Research, Kyoto University
"Games between Males and Females: Mate Guarding, Parental Care, Speciation"
Question - and - Answer
Lecture Tatsuyoshi Saijo
Professor, Institute of Social and Economic Research, Osaka University
"Emergence of Cooperation"
Question - and - Answer
Lecture Tetsukazu Yahara
Professor, Department of Biology, Faculty of Sciences, Kyushu University
"Short-term and Long-term Advantages of Sexual Reproduction: Evidence from Empirical Studies in Asteraceae"
Question - and - Answer
Lecture Ichizo Kobayashi
Assistant Professor, The Institute of Medical Science, The University of Tokyo
"Sex as a Strategy against Selfish Genetic Elements Lessons from Microbial Worlds"
Question - and - Answer
Closing Yoh Iwasa