W. D. Hamilton

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W. D. Hamilton
W D Hamilton.jpg
W. D. Hamilton, 1996
William Donald Hamilton

(1936-08-01)1 August 1936
Died7 March 2000(2000-03-07) (aged 63)
Nationality British
Alma mater University College London
London School of Economics
St. John's College, Cambridge
Known for Kin selection, Hamilton's rule
Awards Newcomb Cleveland Prize (1981)
Linnean Medal (1989)
Kyoto Prize (1993)
Crafoord Prize (1993)
Sewall Wright Award (1998)
Scientific career
Fields Evolutionary biology
Academic advisors John Hajnal
Cedric Smith
Doctoral students Laurence Hurst
Olivia Judson
Influences Ronald Fisher
Influenced Richard Dawkins

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. [1] [2]


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 the gene-centered view of evolution. He is considered one of the forerunners of sociobiology. 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.

Early life

Hamilton was born in 1936 in Cairo, Egypt, the second of seven children. His parents were from New Zealand; his father A. M. Hamilton was an engineer, and his mother B. M. Hamilton was 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 spent 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 Smythe House. As a 12-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 in 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, 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 [ citation needed ]. 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.

Hamilton's rule

Hamilton enrolled in an MSc course in demography at the London School 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 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. 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 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. r is a number between 0 and 1. His two 1964 papers entitled The Genetical Evolution of Social Behavior are now widely referenced. [3]

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 Theoretical 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 reproducing themselves.

Spiteful behaviour

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. [4] 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. [5]

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.

Chasing the Red Queen

Hamilton was an early proponent of the Red Queen theory of the evolution of sex [6] (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 Behaviour.

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.[ citation needed ]

Hamilton also proposed the coevolution theory of autumn leaf color as an example of evolutionary signalling theory. [7]

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, Hamilton and two others ventured on a field trip to the war-torn Democratic Republic of the Congo.[ citation needed ]


He returned to London from Africa on 29 January 2000. He was admitted to University College Hospital, London, on 30 January 2000. He was transferred to 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, the 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. [8]

A secular memorial service (he was an agnostic [9] ) was held at the chapel of 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: [10]

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 this great 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 2005.




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.

Significant papers


  1. "Obituary by Richard Dawkins", The Independent, 10 March 2000. See also his eulogy by Richard Dawkins reprinted in his book A Devil's Chaplain (2003).
  2. BBC Radio 4 – Great Lives – 2 Feb 2010
  3. 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
  4. Brown, Andrew (2000). The Darwin Wars: The Scientific Battle for the Soul of Man. London: Touchstone. ISBN   978-0-684-85145-7.
  5. The Red Queen Hypothesis at Indiana University. Quote: "W. D. Hamilton and John Jaenike were among the earliest pioneers of the idea."
  6. 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 . PMID   11454293.
  7. "ECU Ruling: Great Lives, BBC Radio 4, 2 February 2010". BBC. Retrieved 24 June 2011.
  8. Ullica Segerstrale (28 February 2013). Nature's Oracle: The Life and Work of W.D.Hamilton. OUP Oxford. pp. 383–. ISBN   978-0-19-164277-7.
  9. Hamilton, W. D. (2000). "My intended burial and why". Ethology Ecology and Evolution. 12 (2): 111–122. doi:10.1080/08927014.2000.9522807.[ permanent dead link ]
  10. 1 2 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.

Related Research Articles

An evolutionarily stable strategy (ESS) is a strategy which, if adopted by a population in a given environment, is impenetrable, meaning that it cannot be invaded by any alternative strategy that are initially rare. It is relevant in game theory, behavioural ecology, and evolutionary psychology. An ESS is an equilibrium refinement of the Nash equilibrium. It is a Nash equilibrium that is "evolutionarily" stable: once it is fixed in a population, natural selection alone is sufficient to prevent alternative (mutant) strategies from invading successfully. The theory is not intended to deal with the possibility of gross external changes to the environment that bring new selective forces to bear.

<i>The Selfish Gene</i> 1976 book by Richard Dawkins

The Selfish Gene is a 1976 book on evolution by the biologist Richard Dawkins, in which the author builds upon the principal theory of George C. Williams's Adaptation and Natural Selection (1966). Dawkins uses the term "selfish gene" as a way of expressing the gene-centred view of evolution, popularising ideas developed during the 1960s by W. D. Hamilton and others. From the gene-centred view, it follows that the more two individuals are genetically related, the more sense it makes for them to behave selflessly with each other.

Selfish genetic elements are genetic segments that can enhance their own transmission at the expense of other genes in the genome, even if this has no positive or a net negative effect on organismal fitness. Genomes have traditionally been viewed as cohesive units, with genes acting together to improve the fitness of the organism. However, when genes have some control over their own transmission, the rules can change, and so just like all social groups, genomes are vulnerable to selfish behaviour by their parts.

Kin selection The evolutionary strategy that favours the reproductive success of an organisms relatives, even at a cost to the organisms own survival and reproduction

Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution is driven by kin selection. Kin selection is an instance of inclusive fitness, which combines the number of offspring produced with the number an individual can ensure the production of by supporting others, such as siblings.

Modern synthesis (20th century) Combination of Darwins theory of evolution with natural selection and Mendels findings on heredity

The Modern Synthesis was the early 20th-century synthesis reconciling Charles Darwin's theory of evolution and Gregor Mendel's ideas on heredity in a joint mathematical framework. Julian Huxley coined the term in his 1942 book, Evolution: The Modern Synthesis.

Group selection Proposed mechanism of evolution

Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the more conventional level of the individual.

Fisher's fundamental theorem of natural selection is an idea about genetic variance in population genetics developed by the statistician and evolutionary biologist Ronald Fisher. The proper way of applying the abstract mathematics of the theorem to actual biology has been a matter of some debate.

George C. Williams (biologist) evolutionary biologist from the United States

George Christopher Williams was an American evolutionary biologist.

<i>Journal of Theoretical Biology</i> Academic journal

The Journal of Theoretical Biology is a biweekly peer-reviewed scientific journal covering theoretical biology, as well as mathematical, computational and statistical aspects of biology. Some research areas covered by the journal include cell biology, evolutionary biology, population genetics, morphogenesis, and immunology. It is often referred to informally by the initials 'JTB'.

George R. Price American mathematician

George Robert Price was an American population geneticist.

Index of evolutionary biology articles Wikimedia list article

This is a list of topics in evolutionary biology.

Inclusive fitness A measure of evolutionary success based on the number of offspring the individual supports

In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964:

Evolutionary game theory (EGT) is the application of game theory to evolving populations in biology. It defines a framework of contests, strategies, and analytics into which Darwinian competition can be modelled. It originated in 1973 with John Maynard Smith and George R. Price's formalisation of contests, analysed as strategies, and the mathematical criteria that can be used to predict the results of competing strategies.

"The Genetical Evolution of Social Behaviour" is a 1964 scientific paper by the British evolutionary biologist W.D. Hamilton in which he mathematically lays out the basis for inclusive fitness.

Gene-centered view of evolution Reasoning that since heritable information is passed from generation to generation almost exclusively by DNA, natural selection and evolution are best considered from the perspective of genes

The gene-centered view of evolution, gene's eye view, gene selection theory, or selfish gene theory holds that adaptive evolution occurs through the differential survival of competing genes, increasing the allele frequency of those alleles whose phenotypic trait effects successfully promote their own propagation, with gene defined as "not just one single physical bit of DNA [but] all replicas of a particular bit of DNA distributed throughout the world". The proponents of this viewpoint argue that, since heritable information is passed from generation to generation almost exclusively by DNA, natural selection and evolution are best considered from the perspective of genes.

In biology, the idea of an unbeatable strategy was proposed by W.D. Hamilton in his 1967 paper on sex ratios in Science. In this paper Hamilton discusses sex ratios as strategies in a game, and cites Verner as using this language in his 1965 paper which "claims to show that, given factors causing fluctuations of the population's primary sex ratio, a 1:1 sex-ratio production proves the best overall genotypic strategy".

In biology, altruism refers to behaviour by an individual that increases the fitness of another individual while decreasing the fitness of the actor. Altruism in this sense is different from the philosophical concept of altruism, in which an action would only be called "altruistic" if it was done with the conscious intention of helping another. In the behavioural sense, there is no such requirement. As such, it is not evaluated in moral terms—it is the consequences of an action for reproductive fitness that determine whether the action is considered altruistic, not the intentions, if any, with which the action is performed.

Intragenomic conflict refers to the evolutionary phenomenon where genes have phenotypic effects that promote their own transmission in detriment of the transmission of other genes that reside in the same genome. The selfish gene theory postulates that natural selection will increase the frequency of those genes whose phenotypic effects cause their transmission to new organisms, and most genes achieve this by cooperating with other genes in the same genome to build an organism capable of reproducing and/or helping kin to reproduce. The assumption of the prevalence of intragenomic cooperation underlies the organism-centered concept of inclusive fitness. However, conflict among genes in the same genome may arise both in events related to reproduction and altruism.

Fisher's principle is an evolutionary model that explains why the sex ratio of most species that produce offspring through sexual reproduction is approximately 1:1 between males and females. A. W. F. Edwards has remarked that it is "probably the most celebrated argument in evolutionary biology".

Darwinian anthropology describes an approach to anthropological analysis which employs various theories from Darwinian evolutionary biology. Whilst there are a number of areas of research that can come under this broad description some specific research projects have been closely associated with the label. A prominent example is the project that developed in the mid 1970s with the goal of applying sociobiological perspectives to explain patterns of human social relationships, particularly kinship patterns across human cultures.