William Donald Hamilton, FRS (1 August 1936 – 7 March 2000) was an
English evolutionary biologist, widely recognised as one of the most
significant evolutionary theorists of the 20th century.
Hamilton became famous through his theoretical work expounding a
rigorous genetic basis for the existence of altruism, an insight that
was a key part of the development of a gene-centric view of evolution.
He is considered one of the forerunners of sociobiology, as
popularized by E. O. Wilson. Hamilton also published important
work on sex ratios and the evolution of sex. From 1984 to his death in
2000, he was a
Royal Society Research Professor at Oxford University.
1 Early life
2 Hamilton's rule
3 Spiteful behaviour
4 Extraordinary sex ratios
5 Chasing the Red Queen
6 Return to Britain
7 Social evolution
8 Expedition to the Congo
12.1 Collected papers
12.2 Significant papers
15 External links
Hamilton was born in 1936 in Cairo, Egypt, the second of seven
children. His parents were from New Zealand; his father A. M. Hamilton
an engineer, and his mother B. M. Hamilton a medical doctor. The
Hamilton family settled in Kent. During the Second World War, the
young Hamilton 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.
New Naturalist book Butterflies, which introduced him to the
principles of evolution by natural selection, genetics, and population
He was educated at Tonbridge School, where he was in Smythe House. As
a twelve-year-old he was seriously injured while playing with
explosives his father had. These were left over from his father making
hand grenades for the Home Guard during World War II; he had to
have a thoracotomy and fingers on his right hand had to be amputated
King's College Hospital
King's College Hospital to save his life, and he was left with
scarring and needed six months to recover.
Before going up to the
University of Cambridge
University of Cambridge he travelled in France
and completed two years of national service. As an undergraduate at
St. John's College, he was uninspired by the "many biologists [who]
hardly seemed to believe in evolution". He was intrigued by Ronald
Fisher's book The Genetical Theory of Natural Selection; but Fisher
lacked standing at Cambridge, being viewed as only a
statistician. Hamilton was excited by Fisher's
chapters on eugenics. In earlier chapters, Fisher provided a
mathematical basis for the genetics of evolution and Hamilton later
blamed Fisher's book for his getting only a 2:1 degree.
Main article: Hamilton's rule
Hamilton enrolled in an MSc course in demography at the
of Economics (LSE), under Norman Carrier, who helped secure various
grants for his studies. Later, when his work became more mathematical
and genetical, he had his supervision transferred to
John Hajnal of
the LSE and Cedric Smith of University College
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. This
became Hamilton's rule: in each behaviour-evoking situation, the
individual assesses his neighbour's fitness against his own according
to the coefficients of relationship appropriate to the situation.
Algebraically, the rule posits that a costly action should be
displaystyle C<rtimes B
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. r
is a number between 0 and 1. His two 1964 papers entitled The
Evolution of Social Behavior are now widely referenced.
The proof and discussion of its consequences, however, involved
detailed mathematics, and two reviewers passed over the paper. The
third, John Maynard Smith, did not completely understand it either,
but recognised its significance. Having his work passed over later led
to friction between Hamilton and Maynard Smith, as Hamilton thought
Smith had held his work back to claim credit for the idea (during the
review period Maynard Smith published a paper that referred briefly to
similar ideas). The Hamilton paper was printed in the Journal of
Biology and, when first published, was largely ignored.
Recognition of its significance gradually increased to the point that
it is now routinely cited in biology books.
Much of the discussion relates 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
Main article: Hamiltonian spite
In his 1970 paper Selfish and Spiteful Behaviour in an Evolutionary
Model Hamilton considers the question of whether harm inflicted upon
an organism must inevitably be a byproduct of adaptations for
survival. What of possible cases where an organism is deliberately
harming others without apparent benefit to the self? Such behaviour
Hamilton calls spiteful. It can be explained as the increase in the
chance of an organism's genetic alleles to be passed to the next
generations by harming those that are less closely related than
relationship by chance.
Spite, however, is unlikely ever to be elaborated into any complex
forms of adaptation. Targets of aggression are likely to act in
revenge, and the majority of pairs of individuals (assuming a
panmictic species) exhibit a roughly average level of genetic
relatedness, making the selection of targets of spite problematic.
Extraordinary sex ratios
Between 1964 and 1977 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
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
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
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
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.
Chasing the Red Queen
Hamilton was an early proponent of the Red Queen theory of the
evolution of sex (separate from the other theory of the same name
previously 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.
Return to 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 in the Department of Zoology at Oxford, and a Fellow of New
College, where he remained until his death.
From 1994 Hamilton found companionship with Maria Luisa Bozzi, an
Italian science journalist and author.
His collected papers, entitled Narrow Roads of Gene Land, began to be
published in 1996. The first volume was entitled
Evolution of Social
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.
Hamilton also proposed the coevolution theory of autumn leaf color as
an example of evolutionary signalling theory.
Expedition to the Congo
During the 1990s Hamilton became increasingly interested in the
controversial argument that the origin of HIV lay in oral polio
vaccines trials conducted by
Hilary Koprowski in Africa during the
1950s. Letters by Hamilton on the topic to the major peer-reviewed
journals were rejected. To look for indirect evidence of the OPV
hypothesis by assessing natural levels of simian immunodeficiency
virus, in primates, in early 2000 he and two others ventured on a
field trip to the war-torn Democratic Republic of the Congo.
He returned to
London from Africa on 29 January 2000. He was admitted
to University College Hospital, London, on 30 January 2000. He was
Middlesex Hospital on 5 February 2000 and died there on
7 March 2000. An inquest was held on 10 May 2000 at Westminster
Coroner's Court to inquire into rumours about the cause of his death.
The coroner concluded that his death was due to "Multi-organ failure
due to upper gastrointestinal haemorrhage due to a duodenal
diverticulum and arterial bleed through a mucosal ulcer". Following
reports attributing his death to complications arising from malaria,
BBC Editorial Complaints Unit's investigation established that he
had contracted malaria during his final African expedition. However,
the pathologist had suggested the possibility that the ulceration and
consequent haemorrhage had resulted from a pill (which might have been
taken because of malarial symptoms) lodging in the diverticulum; but,
even if this suggestion were correct, the link between malaria and the
observed causes of death would be entirely indirect.
A secular memorial service (he was an agnostic) was held at the
New College, Oxford
New College, Oxford on 1 July 2000, organised by Richard
Dawkins. He was buried near
Wytham Woods. He, however, had written an
essay on My intended burial and why in which he wrote:
I will leave a sum in my last will for my body to be carried to Brazil
and to these forests. It will be laid out in a manner secure against
the possums and the vultures just as we make our chickens secure; and
Coprophanaeus beetle will bury me. They will enter, will
bury, will live on my flesh; and in the shape of their children and
mine, I will escape death. No worm for me nor sordid fly, I will buzz
in the dusk like a huge bumble bee. I will be many, buzz even as a
swarm of motorbikes, be borne, body by flying body out into the
Brazilian wilderness beneath the stars, lofted under those beautiful
and un-fused elytra which we will all hold over our backs. So finally
I too will shine like a violet ground beetle under a stone.
The second volume of his collected papers,
Evolution of Sex, was
published in 2002, and the third and final volume, Last Words, in
1978 Foreign Honorary Member of American Academy of Arts and Sciences
1980 Fellow of the
Royal Society of London
Newcomb Cleveland Prize of the American Association for the
Advancement of Science
Darwin Medal of the
Royal Society of London
1989 Scientific Medal of the Linnean Society
Frink Medal of Zoological Society of London
1992/3 Wander Prize of the University of Bern
Crafoord Prize of the Royal Swedish Academy of Sciences
Kyoto Prize of the Inamori Foundation
1995 Fyssen Prize of the Fyssen Foundation
1997 Honorary title of Academician of Science in Finland
Alan Grafen has written a biographical memoir for the Royal
A biographical book has also been published by Ullica
Segerstråle : Segerstråle, U. 2013. Nature's oracle: the life
and work of W. D. Hamilton.
Oxford University Press.
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:
Oxford University Press, Oxford.
Hamilton W.D. (2002) Narrow Roads of Gene Land vol. 2:
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
Hamilton, W. (1964). "The genetical evolution of social behaviour. I".
Journal of Theoretical Biology. 7 (1): 1–16.
doi:10.1016/0022-5193(64)90038-4. PMID 5875341.
Hamilton, W. (1964). "The genetical evolution of social behaviour.
II". Journal of Theoretical Biology. 7 (1): 17–52.
doi:10.1016/0022-5193(64)90039-6. PMID 5875340.
Hamilton, W. (1966). "The moulding of senescence by natural
selection". Journal of Theoretical Biology. 12 (1): 12–45.
doi:10.1016/0022-5193(66)90184-6. PMID 6015424.
Hamilton, W. (1967). "Extraordinary sex ratios. A sex-ratio theory for
sex linkage and inbreeding has new implications in cytogenetics and
entomology". Science. 156 (774): 477–488.
doi:10.1126/science.156.3774.477. PMID 6021675.
Hamilton, W. (1971). "Geometry for the selfish herd". Journal of
Theoretical Biology. 31 (2): 295–311.
doi:10.1016/0022-5193(71)90189-5. PMID 5104951.
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.; Hamilton, W. (1981). "The evolution of cooperation".
Science. 211 (4489): 1390–1396. doi:10.1126/science.7466396.
PMID 7466396. with Robert Axelrod
Hamilton, W.; Zuk, M. (1982). "Heritable true fitness and bright
birds: A role for parasites?". Science. 218 (4570): 384–387.
doi:10.1126/science.7123238. PMID 7123238.
^ "Obituary by Richard Dawkins", The Independent, 10 March 2000. See
also his eulogy by
Richard Dawkins reprinted in his book A Devil's
BBC Radio 4 - Great Lives - 2 Feb 2010
^ Aaen-Stockdale, C. (2017), "Selfish Memes: An Update of Richard
Dawkins' Bibliometric Analysis of Key Papers in Sociobiology",
Publications, 5 (2): 12, doi:10.3390/publications5020012
^ Brown, Andrew (2000). The Darwin Wars: The Scientific Battle for the
Soul of Man. London: Touchstone. ISBN 0-684-85145-8.
^ The Red Queen Hypothesis at Indiana University. Quote: "W. D.
Hamilton and John Jaenike were among the earliest pioneers of the
^ Hamilton, WD; Brown, SP (July 2001). "Autumn tree colours as a
handicap signal". Proc. R. Soc. B. 268 (1475): 1489–1493.
doi:10.1098/rspb.2001.1672. ISSN 0962-8452. PMC 1088768 .
^ "ECU Ruling: Great Lives,
BBC Radio 4, 2 February 2010". BBC.
Retrieved 24 June 2011.
^ Ullica Segerstrale (28 February 2013). Nature's Oracle: The Life and
Work of W.D.Hamilton. OUP Oxford. pp. 383–.
^ Hamilton, W. D. (2000). "My intended burial and why". Ethology
Ecology and Evolution. 12: 111–122.
^ a b Grafen, A. (2004). "William Donald Hamilton. 1 August 1936 -- 7
March 2000" (PDF). Biographical Memoirs of Fellows of the Royal
Society. 50: 109–132. doi:10.1098/rsbm.2004.0009.
Edwards, A. W. F. (1998) Notes and Comments. Edwards, A. W.
F. (1998). "Natural Selection and the Sex Ratio: Fisher's Sources".
The American Naturalist. 151 (6): 564–569. doi:10.1086/286141.
Fisher R. A. (1930). The Genetical Theory of Natural Selection.
Clarendon Press, Oxford.
Ford, E. B. (1945) New Naturalist 1: Butterflies. Collins:
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., Robin Dunbar,
and J. M. Richardson and D. McFarland. (2006) "Kinship and
altruism: a cross-cultural experimental study". British Journal of
Obituaries and reminiscences
Royal Society citation
Truth and Science: Bill Hamilton's legacy
Centro Itinerante de Educação Ambiental e Científica 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
London Review of Books book review
W. D. Hamilton's work in game theory
Evolutionarily stable strategy
Darwinian literary studies
Evolution of emotion
W. D. Hamilton
Carel van Schaik
David Sloan Wilson
E. O. Wilson
George C. Williams
Jerome H. Barkow
Dominic D. P. Johnson
Justin L. Barrett
David F. Bjorklund
David C. Geary
Judith Rich Harris
Aurelio José Figueredo
Douglas T. Kenrick
Simon M. Kirby
Michael T. McGuire
Randolph M. Nesse
David P. Schmitt
Todd K. Shackelford
Peter K. Smith
Mark van Vugt
Center for Evolutionary Psychology
Human Behavior and
Max Planck Institute for Human Cognitive and Brain Sciences
New England Complex Systems Institute
The Adapted Mind
Evolution of Human Sexuality
Evolution and Human Behavior
Evolutionary psychology and culture
Criticism of evolutionary psychology
Evolutionary psychology research groups and centers
Bibliography of evolution and human behavior
Evolutionary biology portal
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