Extraordinary Sex Ratios
Between 1964 and 1978 Hamilton was a lecturer at University College London. Whilst there he published a paper in Science on "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary" sex ratios were nearly always 1:1 (but see Edwards 1998), and likewise extraordinary sex ratios, particularly in wasps, needed explanations. Hamilton had been introduced to the idea and formulated its solution in 1960 when he had been assigned to help Fisher's pupil A.W.F. Edwards test the Fisherian sex ratio hypothesis. Hamilton combined his extensive knowledge of natural history with deep insight into the problem, opening up a whole new area of research.
The paper was also notable for introducing the concept of the "unbeatable strategy", which John Maynard Smith and George R. Price were to develop into the evolutionarily stable strategy (ESS), a concept in game theory not limited to evolutionary biology. Price had originally come to Hamilton after deriving the Price equation, and thus rederiving Hamilton's rule. Maynard Smith later peer reviewed one of Price's papers, and drew inspiration from it. The paper was not published but Maynard Smith offered to make Price a co-author of his ESS paper, which helped to improve relations between the men. Price committed suicide in 1975, and Hamilton and Maynard Smith were among the few present at the funeral.
Hamilton was regarded as a poor lecturer. This shortcoming would not affect the popularity of his work, however, as it was popularised by Richard Dawkins in Dawkins' 1976 book The Selfish Gene.
In 1966 he married Christine Friess and they were to have three daughters, Helen, Ruth and Rowena. 26 years later they amicably separated.
Hamilton was a visiting professor at Harvard University and later spent nine months with the Royal Society's and the Royal Geographical Society's Xavantina-Cachimbo Expedition as a visiting professor at the University of São Paulo.
From 1978 Hamilton was Professor of Evolutionary Biology at the University of Michigan. Simultaneously, he was elected a Foreign Honorary Member of American Academy of Arts and Sciences. His arrival sparked protests and sit-ins from students who did not like his association with sociobiology. There he worked with the political scientist Robert Axelrod on the prisoner's dilemma, and was a member of the BACH group with original members Arthur Burks, Robert Axelrod, Michael Cohen and John Holland.
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