W. D. Hamilton, 1996
'William Donald Hamilton',
F.R.S. (
1 August 1936 —
7 March 2000) was a
British evolutionary biologist, considered one of the greatest evolutionary theorists of the 20th century.
[1] From 1984 to his death in 2000, he was the Royal Society Research Professor at
Oxford University. Hamilton became famous for his theoretical work expounding a rigorous genetic basis for the existence of
kin selection. This insight was a key part of the development of a
gene-centric view of evolution, and he can therefore be seen as one of the forerunners of the discipline of
sociobiology popularized by
Edward Osborne Wilson. Hamilton also published important work on
sex ratios and the
evolution of sex.
Biography
Early life
Hamilton was born in 1936 in
Cairo,
Egypt, the second eldest of six children. His father,
A. M. Hamilton was a
New Zealand-born engineer, and his mother, B. M. Hamilton was a
medical doctor.
The Hamilton family moved to Kent when Bill was young and during the
Second World War he was evacuated to
Edinburgh. He had an interest in natural history from an early age and would spend his spare time collecting
butterflies and other insects. In 1946 he discovered
E.B. Ford's ''
New Naturalist'' book ''Butterflies'', which introduced him to the principles of
evolution by
natural selection,
genetics and
population genetics.
He was educated at
Tonbridge School, where he was in the School House. As a 12-year old he was seriously injured while playing with
explosives his father had left over from when he made
hand grenades for the
Home Guard during the
Second World War, an accident that probably would have killed him had his mother not been medically qualified. A
thoracotomy in
King’s College Hospital saved his life, but the explosion left him with amputated fingers on his right hand and scarring on his body — he took six months to recover.
Hamilton stayed on an extra term at Tonbridge in order to complete the
Cambridge entrance examinations, and then travelled in
France. He then completed two years of
national service. As an
undergraduate at
St. John's College, he was uninspired by the fact that there "many biologists hardly seemed to believe in evolution". Nevertheless, he came across
Ronald Fisher's book ''
The Genetical Theory of Natural Selection''; Fisher lacked standing at Cambridge as he was viewed only as a statistician. Hamilton wrote on a postcard to his sister Mary on the day he found the book excited by its chapters on
eugenics. In earlier chapters, Fisher provided a mathematical basis for the genetics of
evolution. Working through the stodgy prose, Hamilton later blamed Fisher's book for getting only a
2:1 degree.
===
Hamilton's rule===
Hamilton having various ideas and problems enrolled on an MSc course in human demographics at the
London School of Economics (LSE), under
Norman Carrier who secured for him various grants. Later when work became more mathematical and then genetical, he had his supervision transferred to
John Hajnal of the LSE and
Cedric Smith of
University College London (UCL).
Both Fisher and
J. B. S. Haldane had seen a problem in how organisms could increase the fitness of their own genes by aiding their close relatives, but not recognised its significance or properly formulated it. Hamilton worked through several examples, and eventually realised that the number that kept falling out of his calculations was
Sewall Wright's
coefficient of relationship. Thus became Hamilton's rule. Briefly, the rule is that a costly action should be performed if:
:
Where ''C'' is the cost in fitness to the actor, ''R'' the genetic relatedness between the actor and the recipient and ''B'' is the fitness benefit to the recipient. Fitness costs and benefits are measured in fecundity. His two 1964 papers entitled ''
The Genetical Evolution of Social Behavior'' are now universally referenced.
The proof and discussion of its consequences however involved heavy mathematics, and was passed over by two reviewers. The third,
John Maynard Smith, did not completely understand it either, but recognised its significance; this passing over would later lead to friction between Hamilton and Maynard Smith, Hamilton feeling that Maynard Smith had held his work back to claim credit for the idea himself. The paper was printed in the relatively obscure ''
Journal of Theoretical Biology'', and when first published was largely ignored. The significance of it gradually increased, to the point where they are routinely cited in biology books. To date, however, there have been no empirical studies that have calculated values for R, B, and C to determine if Hamilton's rule is ever satisfied in nature; as such, even after more than 40 years, the theory remains unproven, though predictions based upon the theory are largely supported.
A large part of the discussion related to the evolution of
eusociality in insects of the order
Hymenoptera (
ants,
bees and
wasps) based on their unusual
haplodiploid sex-determination system. This system means that females are more closely related to their sisters than to their own (potential) offspring. Thus, Hamilton reasoned, a "costly action" would be better spent in helping to raise their sisters, rather than reproducing themselves.
Extraordinary sex ratios
Between 1964 and 1978 Hamilton was a lecturer at
Imperial 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 Geographic 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
economist Robert Axelrod on the
prisoner's dilemma.
Chasing the Red Queen
Hamilton was an early proponent of the
Red Queen theory of the
evolution of sex,
[1] first proposed by
Leigh Van Valen. This was named for a character in
Lewis Carroll's ''
Through the Looking Glass'', who is continuously running but never actually travels any distance:
:''"Well, in our country," said Alice, still panting a little, "you'd generally get to somewhere else—if you ran very fast for a long time, as we've been doing."''
:''"A slow sort of country!" said the Queen. "Now, here, you see, it takes all the running you can do, to keep in the same place. If you want to get somewhere else, you must run at least twice as fast as that!"''
(Carroll, pp. 46)
This theory hypothesizes that sex evolved because new and unfamiliar combinations of genes could be presented to
parasites, preventing the parasite from preying on that organism—species with sex were able to continuously "run away" from their parasites. Likewise, parasites were able to evolve mechanisms to get around the organism's new set of genes, thus perpetuating an endless race.
Back in Britain
In 1980 he was elected a
Fellow of the Royal Society, and in 1984 he was invited by
Richard Southwood to be the Royal Society Research Professor at
New College,
Oxford, Department of Zoology, where he remained until his death.
From 1994 Hamilton found companionship with Maria Luisa Bozzi, an
Italian science
journalist and author.
On the Origin of HIV
During the 1990s Hamilton became increasingly convinced by the controversial argument that the origin of the
HIV virus lay in oral
polio vaccines (the
OPV AIDS hypothesis) in Africa during the 1950s. Letters by Hamilton to ''
Science'' were rejected by the journal, amid accusations that the medical establishment were ranging against the OPV hypothesis.
To find indirect evidence of the OPV hypothesis by assessing natural levels of SIV in primates, he and two others ventured on a field trip to the war-torn
Democratic Republic of the Congo, where he contracted
malaria. He was rushed home and spent six weeks in hospital before dying from a cerebral haemorrhage.
Postscript
A secular memorial service (he was an
atheist) was held at the Chapel of
New College, Oxford on Saturday
1 July 2000, organised by
Richard Dawkins.
His body was interred in
Wytham Woods. He however had written an essay on ''My intented burial and why'' in which he wrote:
The second volume of his collected papers was published in 2002.
Social evolution
The field of
social evolution, of which
Hamilton's rule has central importance, is broadly defined as being the study of the evolution of social behaviours, i.e. those that impact on the fitness of individuals other than the actor. Social behaviours can be categorized according to the fitness consequences they entail for the actor and recipient. A behaviour that increases the direct fitness of the actor is mutually beneficial if the recipient also benefits, and selfish if the recipient suffers a loss. A behaviour that reduces the fitness of the actor is altruistic if the recipient benefits, and spiteful if the recipient suffers a loss. This classification was first proposed by Hamilton in 1964.
Awards
★ 1978 Foreign Honorary Member of
American Academy of Arts and Sciences
★ 1980 Fellow of the
Royal Society of London
★ 1982
Newcomb Cleveland Prize of the
American Association for the Advancement of Science
★ 1988
Darwin Medal of the
Royal Society of London
★ 1989 Scientific Medal of the
Linnean Society
★ 1991
Frink Medal of Zoological Society of London
★ 1992/3
Wander Prize of the
University of Bern
★ 1993
Crafoord Prize of the
Royal Swedish Academy of Sciences
★ 1993
Kyoto Prize of the
Inamori Foundation
★ 1995
Frissen Prize of the
Fyssen Foundation
Biographies
★
Alan Grafen has written a biographical memoir for the Royal Society. See http://users.ox.ac.uk/~grafen/cv/WDH_memoir.pdf
★ A book is also in press: SegerstrÃ¥le, U. 2007 ''Nature's oracle: an intellectual biography of evolutionist W. D. Hamilton.'' Oxford University Press. See http://www.oup.com/uk/catalogue/?ci=9780198607274
Works
Collected papers
Hamilton started to publish his collected papers starting in 1996, along the lines of Fisher's collected papers, with short essays giving each paper context. He died after the preparation of the second volume, so the essays for the third volume come from his coauthors.
★ 'Hamilton, W.D.' (1996) ''Narrow Roads of Gene Land vol. 1: Evolution of Social Behaviour'' Oxford University Press,Oxford. ISBN 0-7167-4530-5
★ 'Hamilton, W.D.' (2002) ''Narrow Roads of Gene Land vol. 2: Evolution of Sex'' Oxford University Press,Oxford. ISBN 0-19-850336-9
★ 'Hamilton, W.D.' (2005) ''Narrow roads of Gene Land, vol. 3: Last Words'' (with essays by coauthors, ed. M. Ridley). Oxford University Press, Oxford. ISBN 0-19-856690-5
Significant papers by Hamilton
★ 'Hamilton, W.D.' (1964). The genetical evolution of social behaviour I and II. — ''Journal of Theoretical Biology'' '7': 1-16 and 17-52.
pubmed I pubmed II
★ 'Hamilton, W.D.' (1967). Extraordinary sex ratios. ''
Science'' '156': 477-488.
pubmed JSTOR
★ 'Hamilton, W.D.' (1975).
Innate social aptitudes of man: an approach from evolutionary genetics. in R. Fox (ed.), ''Biosocial Anthropology, Malaby Press, London'', 133-53.
★ '
Axelrod, R. and W.D. Hamilton' (1981) The evolution of co-operation ''
Science'' '211': 1390-6
Pubmed,
JSTOR
★ 'Hamilton, W.D.' (2000) My intended burial and why, ''Ethology Ecology and Evolution'' '12' 111-122
link
Other references
★ '
Edwards, A.W.F.' (1998), Notes and Comments. Natural selection and sex ratio: Fisher's sources. ''American Naturalist'' '151': 564-569
★ '
Fisher, R.A.' (1930). ''
The Genetical Theory of Natural Selection''. Clarendon Press, Oxford.
★ '
Ford E.B.' (1945) New Naturalist 1: Butterflies. Collins: London.
★ '
Maynard Smith, J. and
G.R. Price' (1973) The logic of animal conflict. ''
Nature'' '146': 15—18.
★ '
Dawkins, R.' (1989),
The Selfish Gene, 2nd ed. Oxford University Press.
★ 'Madsen, E.A., Tunney, R., Fieldman, G.,
Plotkin, H.C.,
Dunbar, R.I.M., Richardson, J.M., & McFarland, D.' (2006) Kinship and altruism: A cross-cultural experimental study. ''British Journal of Psychology''
[2]
References
1. The Red Queen Hypothesis at Indiana University. Quote: ''"W.D. Hamilton and John Jaenike were among the earliest pioneers of the idea."''
★
Obituaries and reminiscences
★
Royal Society citation
★
Truth and Science: Bill HamiltonÃs legacy
★
Centro Itinerante de Educação Ambiental e CientÃfica Bill Hamilton (The Bill Hamilton Itinerant Centre for Environmental and Scientific Education) (in Portuguese)
★
Non-mathematical excerpts from Hamilton 1964
★
"If you have a simple idea, state it simply" a 1996 interview with Hamilton
Game Theory
★
W.D. Hamilton's work in game theory
AIDS
★
Polio vaccines and AIDS
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