William D. Hamilton

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William Donald "Bill" Hamilton (born August 1, 1936 in Cairo , Egypt , † March 7, 2000 in London ) was a British biologist who carried out research in the fields of theoretical biology , ethology , evolutionary biology , zoologist and genetics . He became famous for his theoretical work that the genetic basis for the theory of kin selection (kin selection) delivered. It can be seen as a forerunner of the sociobiology established by Edward O. Wilson .


Early years

Hamilton was born in Cairo in 1936, the second oldest of six children. His father, AM Hamilton, was a New Zealand born engineer, and his mother, BM Hamilton, was a medical doctor.

The Hamilton family moved to Kent when Bill was a boy. During the Second World War he was evacuated to Edinburgh . He became interested in natural history at an early age and spent his free time collecting butterflies and other insects. In 1946 he discovered the book Butterflies (Butterflies) by EB Ford , which introduced him to the principles of evolution .

He was educated at Tonbridge School where he lived in the school house. At the age of 12, he was seriously wounded while playing with explosives his father had left when he was making hand grenades for home defense during World War II. His right hand had to be amputated and it took six months to recover.

During his early years at St John's College , Cambridge University , graduating ( BS ) in 1960, he was significantly influenced by Ronald Fisher's book The Genetical Theory of Natural Selection , which provided a mathematical basis for evolutionary genetics. In the main, it went against the notions of group selection .

Hamilton's rule

During his time at University College London and the London School of Economics and Political Science , Hamilton wrote his doctoral thesis in 1968 on the principles later known as 'Hamilton's Rule' of overall fitness . His work on this subject is cited worldwide today.

The overall fitness of a living being can therefore be measured as the number of its own genes that are passed on to the next generation. According to John Maynard Smith , it is composed of

a) direct fitness, one's own genes in one's own offspring, and

b) indirect fitness, one's own genes, which were also passed on to other offspring by relatives.

As relatives some of the same genes have as an individual, this encourages behavior by workers who pass on their own genetic material ( kin selection , kin selection ). This altruism is only successful and spreads if the benefit to the person who shows the altruistic behavior is greater than the cost he has to invest in it (Hamilton's rule).

In mathematical terms, the ratio of benefits (B) to costs (C) must be greater than one divided by the degree of kinship.


with B: benefit; C: cost; r: relatedness

Example: An animal that, through its help, does without two offspring of its own (C = 2), but instead helps a sibling (degree of relationship between siblings in diploid organisms (r = 0.5) to produce five additional offspring (B = 5) , has a higher overall fitness than an animal that "selfishly" does not help.

Taking into account the various degrees of relationship to the recipient and to one's own descendants, the following formula results:

: Degree of relationship of the giver to the descendants of the recipient; : Degree of relationship of the giver to his own offspring

The above formula went a long way toward understanding altruism in social insects. Due to the unusual haplodiploidy of social insects (ants, bees and wasps), full sisters of a nest have a relationship coefficient of 0.75 with one another, with their full brothers 0.25. However, these workers are only related to their own offspring by 50% (r = 0.5), i.e. less than to the sisters. As a result, if the queen has mated only once, it is genetically more beneficial for social insect workers to raise their own sisters as daughters.

Exceptional gender ratios

Between 1964 and 1978, Hamilton was a lecturer at Imperial College London . There he published an article in Science on 'Extraordinary Gender Relations'. In 1930 Ronald A. Fisher proposed a model of why the normal gender ratio is almost always 1: 1 and that unusual ratios such as that of wasps require an explanation. This opened up a whole new area of ​​research. The paper introduced the concept of unbeatable strategy, which John Maynard Smith and George R. Price further developed into the evolutionarily stable strategy ESS, a concept of game theory that was not limited to evolutionary biology.

His work gained popularity when it was introduced by Richard Dawkins in Dawkins' 1976 book The Selfish Gene .

In 1976 he married Christine Friess, they had three daughters, Helen, Ruth and Rowena. They later got divorced.

He was visiting professor at Harvard University and later spent nine months with the Royal Society and the Royal Geographical Society 'Xavantina-Cachimbo Expedition' as visiting professor at the University of São Paulo .

From 1978 on, he was Professor of Evolutionary Biology at the University of Michigan . At the same time he was elected as a foreign honorary member of the American Academy of Arts and Sciences . His arrival sparked protests and sit-in strikes among students who did not share his views on sociobiology .

Back in the UK

In 1980 he was elected a member of the Royal Society and in 1984 he became Royal Society Research Professor at New College, Oxford University , Zoology Department, where he remained until his death.

From 1994 onwards he lived with Maria Luisa Bozzi, an Italian writer.

On the development of AIDS

During the 1990s, Hamilton became increasingly convinced that the origin of the AIDS epidemic in contaminated serum in oral polio vaccines (OPV) was in Africa during the 1950s (the OPV AIDS hypothesis ) . Letters from Hamilton to Science were rejected by the magazine, complaining that the medical establishment would crack down on the OPV AIDS hypothesis.

In order to obtain evidence for the OPV-AIDS hypothesis, one wanted to determine the natural level of simian immunodeficiency virus (SIV) in primates. To do this, Hamilton and two other colleagues ventured into the war-torn Democratic Republic of the Congo , where he contracted malaria . He was brought home and spent seven weeks in the hospital before he died.


A secular memorial service (he was an atheist ) was held on Saturday July 1, 2000 in the Chapel of New College University of Oxford , organized by Richard Dawkins .



  • WD Hamilton (1963) The evolution of altruistic behavior. - The American Naturalist 97: 354-356.
  • WD Hamilton (1964) The genetical evolution of social behavior I and II. - Journal of Theoretical Biology 7: 1-16 and 17-52. PMID 5875341 , PMID 5875340 .
  • WD Hamilton (1966) The molding of senescence by natural selection. - Journal of Theoretical Biology 12: 12-45.
  • WD Hamilton (1967) Extraordinary sex ratios. Science 156: 477-488. PMID 6021675 JSTOR
  • WD Hamilton (1970) Selfish and spiteful behavior in an evolutionary model. - Nature 228: 1218-1220.
  • WD Hamilton (1971) The geometry of the selfish herd. - Journal of Theoretical Biology 31: 295-311.
  • WD Hamilton (1972) Altruism and related phenomena, mainly in social insects. - Annual Review of Ecology and Systematics 3: 193-232.
  • WD Hamilton (1975) Innate social aptitudes of man: an approach from evolutionary genetics. ( Memento of October 9, 2004 in the Internet Archive ) in R. Fox (ed.), Biosocial Anthropology, Malaby Press, London , 133-53.
  • WD Hamilton (1980) Sex versus non-sex versus parasite. - Oikos 35: 282-290.
  • Axelrod, R. and WD Hamilton (1981) The evolution of co-operation Science 211: 1390-6 Pubmed , JSTOR
  • WD Hamilton and Marlene Zuk (1982) Heritable true fitness and bright birds - a role for parasites. Science 218: 384-387.
  • WD Hamilton (1996) Narrow Roads in Gene Land vol. 1 Oxford University Press, Oxford. ISBN 0-7167-4530-5 .
  • WD Hamilton (2000) My intended burial and why, Ethology Ecology and Evolution 12 111-122 link ( Memento of February 8, 2006 in the Internet Archive )
  • WD Hamilton (2002) Narrow Roads in Gene Land vol. 2 Oxford University Press, Oxford. ISBN 0-19-850336-9 .
  • AWF Edwards (1998), Notes and Comments. Natural selection and sex ratio: Fisher's sources. American Naturalist 151: 564-569.
  • Ronald Fisher (1930) The Genetical Theory of Natural Selection . Clarendon Press, Oxford.
  • EB Ford (1945) New Naturalist 1: Butterflies. Collins: London.
  • John Maynard Smith and George R. Price (1973) The logic of animal conflict. Nature 146: 15-18.


  • Ullica Segerstrale: Nature's Oracle: The Life and Work of WD Hamilton. Oxford University Press, USA, 2013, ISBN 978-0-19-860727-4 .

Web links

Individual evidence

  1. ^ Member History: William D. Hamilton. American Philosophical Society, accessed September 20, 2018 .