A Profile of John Maynard Smith: Game Theory and the Evolution of Sex

A Profile of John Maynard Smith: Game Theory and the Evolution of Sex
Page content


A leading figure in the field of genetics and evolutionary biology, the British research theorist John Maynard Smith is known for using game theory to explain animal behavior issues and for introducing the concept of Evolutionary Stable Strategy (ESS). He also worked on -

  • Understanding evolution in microbial organisms
  • Trying to discover a historical commonality in the different complex ways in which different life-forms evolved.
  • Understanding genetic repetition in an organism’s development
  • Assessing DNA recombinational rates

Game Theory

The history of game theory arguments actually goes back to the time of Plato, but the term game theory is a modern one and was originally formulated by Von Neumann and Morgenstern in 1953 to explain various human economic behaviors. Later game theory was applied to understand human psychological, sociological and political behaviors. Ronald Fisher, in the 1930s, used it in regards to animal behavior, and Lewontin, in 1961, was the first to apply it to evolutionary biology. Lewontin, and later Slobodkin and Rapoport in 1974, took the approach of a species versus nature game, the point of which was to find strategies to avoid or minimize the chances of extinction.

John Maynard Smith, in his 1982 book ‘Evolution and the Theory of Games’, took a different line of thought -animals playing games against each other, each individual making choices, like finding a mate, finding food resources, etc., that depended on the choices made by other individuals, and this affecting the population dynamics and equilibrium. The equilibrium, which was maintained by evolutionary forces, was dubbed as the Evolutionarily Stable Strategy (ESS) by Maynard Smith in 1973.

Evolution of Sex

John Maynard Smith pondered over the issue of the evolution of sex and why sexual reproduction prevailed in evolution, in spite of the better efficiency of asexual reproduction. He applied mathematics to explain the two-fold cost of sex. In sexual reproduction, only the females are capable of reproduction, while in asexual reproduction each individual is capable of reproduction. This means that the asexual population will grow at faster rate each generation than the sexual population. Why, in this case, hasn’t asexual assumed ascendancy over sexual reproduction? Researchers have presented many different hypotheses in this matter. One of the reasons may be that asexual offspring are perhaps less fit than sexual offspring, since they inherit the same genotype as their parent and there is no room for genetic diversity as in sexual reproduction.

Personal Life, Education and Career

Maynard Smith (born 6 January 1920,London) spent the first part of his career working as an aeronautical engineer. Although he had developed an early interest in the natural world - after moving to Exmoor at the age of eight after his surgeon father’s death and being given a book on birds by his aunt - he had chosen to study engineering at Trinity College, Cambridge, to avoid being roped into his grandfather’s stockbroker firm. Deemed unfit to serve in the Second World War, he spent the duration of the war testing aircraft stresses and designing military aircraft. In 1947, he decided engineering wasn’t the career for him after all and quit to study zoology under J.B.S. Haldane at University College, London.

It was Haldane’s presence that made him chose University College. Haldane, like Maynard, was a former Etonian, and Maynard had heard much about him - nothing to his credit, with his divorce and communist leanings - while at that institution. Maynard’s perception of Eton had been one of snobbish anti-intellectualism, he hadn’t enjoyed his time there one bit, and the general vilification of Haldane had only intrigued him back then. He read his books in the library and, like him, embraced communism soon after leaving Eton. Now he jumped at the chance to study under him and carried out Drosophila genetics research. Apart from Haldane, his interest in genetics had been evoked by Olaf Stapledon’s book “Last and First Men”.

After getting his B.Sc. degree in Zoology in 1951, he joined University College as a Biology Professor in 1952 and continued working there until 1965; apart from teaching, he carried out research on Drosophila genetics and population genetics.


In 1965, he became the Dean of the School of Biological Sciences at Sussex University, a university he had helped found in 1962. He remained at Sussex University until his retirement in 1985 and then became Professor Emeritus there. He was honored by Sussex University by having the Life Sciences Building named after him, and the European Society for Evolutionary Biology established the John Maynard Smith Prize to ward promising young evolutionary biologists. He received many other awards in the course of his career, including the Royal Society Fellowship (1977), Royal Society Darwin Medal (1986), the Balzan Prize (1991), the Linnean Medal (1995), the Crafoord Prize (1999) and the Kyoto Prize (2001).

He wrote a number of research papers in the course of his scientific career as well as a good many books on science -

  • The Theory of Evolution (1958)
  • Mathematical Ideas in Biology (1968)
  • Models in Ecology (1974)
  • The Evolution of Sex (1978)
  • Evolution and the Theory of Games (1982)
  • Evolutionary Genetics (1989)
  • The Major Transitions of Evolution (1995)
  • Animal Signals (2003)

On the personal front, he married Sheila Matthew shortly after he graduating in engineering, and they had three children. Maynard Smith died of lung cancer on 19 April 2004 at his home in Lewes, East Sussex.


The Evolution of Sex, By John Maynard Smith, Cambridge University Press, 1978

Evolution and the theory of games, By John Maynard Smith, Cambridge University Press, 1982