W. D. Hamilton
Encyclopedia
William Donald Hamilton FRS
(1 August 1936 – 7 March 2000) was a British
evolutionary biologist, widely recognised as one of the greatest evolution
ary theorists of the 20th century.
Hamilton became famous through his theoretical work expounding a rigorous genetic
basis for the existence of kin selection
and 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 ratio
s and the evolution of sex
. From 1984 to his death in 2000, he was a Royal Society
Research Professor at Oxford University.
, Egypt
, the second of seven children. His father A. M. Hamilton
was a New Zealand
-born engineer. His mother B. M. Hamilton was a medical doctor, also from New Zealand.
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. 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 making hand grenade
s for the Home Guard
during World War II
; the accident might have killed him if his mother had not been a doctor. The boy had to have a thoracotomy
in King's College Hospital
to save his life, but fingers on his right hand had to be amputated and he was left with scarring on his body. He needed six months to recover.
Hamilton stayed on an extra term at Tonbridge to complete the Cambridge
entrance examination
s, and then travelled in France
. He 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 as he was 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. Working through the stodgy prose, Hamilton later blamed Fisher's book for his getting only a 2:1 degree.
course in human demographics 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
. His two 1964 papers entitled The Genetical 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 Theoretical Biology
and, when first published, was largely ignored. Recognition of its significance gradually increased to the point, and it is now routinely cited in biology books. Although the theory can be proven mathematically, to date no empirical studies have calculated values for R, B, and C to determine if Hamilton's rule is ever satisfied in nature. After more than 40 years, some scientists claim that the theory remains unconfirmed, although predictions based upon it are largely supported. Because B and C represent gains and losses, it is impossible to compare actual gains and losses with the possible ones that would result from an alternate history. Statisticians, however, argue that such comparisons can be made between similar families in which different versions of history play out.
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.
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.
. Whilst there he published a paper in Science
on "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary" sex ratio
s 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
.
theory of the evolution of sex
, 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:
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.
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.
, 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.
Through his collaboration with Hugh N. Comins and Bob May
on evolutionarily stable dispersal strategies, Hamilton acquired an Erdős number
of 5.
Hamilton also proposed the coevolution theory of autumn leaf color
as an example of evolutionary signalling theory
.
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 SIV in primates, in early 2000 he and two others ventured on a field trip to the war-torn Democratic Republic of the Congo
.
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
. He was buried near Wytham Woods
. He, however, had written an essay on My intended burial and why in which he wrote:
The second volume of his collected papers, Evolution of Sex, was published in 2002, and the third and final volume, Last Words, in 2005.
Game Theory
Royal Society
The Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
(1 August 1936 – 7 March 2000) was a British
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...
evolutionary biologist, widely recognised as one of the greatest evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
ary theorists of the 20th century.
Hamilton became famous through his theoretical work expounding a rigorous genetic
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....
basis for the existence of kin selection
Kin selection
Kin selection refers to apparent strategies in evolution that favor the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Charles Darwin was the first to discuss the concept of group/kin selection...
and altruism
Altruism
Altruism is a concern for the welfare of others. It is a traditional virtue in many cultures, and a core aspect of various religious traditions, though the concept of 'others' toward whom concern should be directed can vary among cultures and religions. Altruism is the opposite of...
, 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
Sociobiology
Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...
, as popularized by E. O. Wilson
E. O. Wilson
Edward Osborne Wilson is an American biologist, researcher , theorist , naturalist and author. His biological specialty is myrmecology, the study of ants....
. Hamilton also published important work on sex ratio
Sex ratio
Sex ratio is the ratio of males to females in a population. The primary sex ratio is the ratio at the time of conception, secondary sex ratio is the ratio at time of birth, and tertiary sex ratio is the ratio of mature organisms....
s and the evolution of sex
Evolution of sex
The evolution of sexual reproduction is currently described by several competing scientific hypotheses. All sexually reproducing organisms derive from a common ancestor which was a single celled eukaryotic species. Many protists reproduce sexually, as do the multicellular plants, animals, and fungi...
. From 1984 to his death in 2000, he was a Royal Society
Royal Society
The Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
Research Professor at Oxford University.
Early life
Hamilton was born in 1936 in CairoCairo
Cairo , is the capital of Egypt and the largest city in the Arab world and Africa, and the 16th largest metropolitan area in the world. Nicknamed "The City of a Thousand Minarets" for its preponderance of Islamic architecture, Cairo has long been a centre of the region's political and cultural life...
, Egypt
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...
, the second of seven children. His father A. M. Hamilton
A. M. Hamilton
Archibald Milne Hamilton was a New Zealand-born civil engineer, notable for building the Hamilton Road through Kurdistan and designing the Callender-Hamilton bridge system. His name is also associated with the Callendar-Hamilton aeroplane shed of the late 1930s.- Early life, marriage and children...
was a New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...
-born engineer. His mother B. M. Hamilton was a medical doctor, also from New Zealand.
The Hamilton family settled in Kent
Kent
Kent is a county in southeast England, and is one of the home counties. It borders East Sussex, Surrey and Greater London and has a defined boundary with Essex in the middle of the Thames Estuary. The ceremonial county boundaries of Kent include the shire county of Kent and the unitary borough of...
. During the Second World War, the young Hamilton was evacuated to Edinburgh
Edinburgh
Edinburgh is the capital city of Scotland, the second largest city in Scotland, and the eighth most populous in the United Kingdom. The City of Edinburgh Council governs one of Scotland's 32 local government council areas. The council area includes urban Edinburgh and a rural area...
. 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
New Naturalist
The New Naturalist Library books are a series published by Collins in the United Kingdom, on a variety of natural history topics relevant to the British Isles...
book Butterflies, which introduced him to the principles of evolution by natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
, genetics
Genetics
Genetics , a discipline of biology, is the science of genes, heredity, and variation in living organisms....
and population genetics
Population genetics
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...
.
He was educated at Tonbridge School
Tonbridge School
Tonbridge School is a British boys' independent school for both boarding and day pupils in Tonbridge, Kent, founded in 1553 by Sir Andrew Judd . It is a member of the Eton Group, and has close links with the Worshipful Company of Skinners, one of the oldest London livery companies...
, 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 making hand grenade
Hand grenade
A hand grenade is any small bomb that can be thrown by hand. Hand grenades are classified into three categories, explosive grenades, chemical and gas grenades. Explosive grenades are the most commonly used in modern warfare, and are designed to detonate after impact or after a set amount of time...
s for the Home Guard
British Home Guard
The Home Guard was a defence organisation of the British Army during the Second World War...
during World War II
World War II
World War II, or the Second World War , was a global conflict lasting from 1939 to 1945, involving most of the world's nations—including all of the great powers—eventually forming two opposing military alliances: the Allies and the Axis...
; the accident might have killed him if his mother had not been a doctor. The boy had to have a thoracotomy
Thoracotomy
Thoracotomy is an incision into the pleural space of the chest. It is performed by a surgeon, and, rarely, by emergency physicians, to gain access to the thoracic organs, most commonly the heart, the lungs, the esophagus or thoracic aorta, or for access to the anterior spine such as is necessary...
in King's College Hospital
King's College Hospital
King's College Hospital is an acute care facility in the London Borough of Lambeth, referred to locally and by staff simply as "King's" or abbreviated internally to "KCH"...
to save his life, but fingers on his right hand had to be amputated and he was left with scarring on his body. He needed six months to recover.
Hamilton stayed on an extra term at Tonbridge to complete the Cambridge
University of Cambridge
The University of Cambridge is a public research university located in Cambridge, United Kingdom. It is the second-oldest university in both the United Kingdom and the English-speaking world , and the seventh-oldest globally...
entrance examination
Entrance examination
An entrance examination is an examination that many educational institutions use to select students for admission. These exams may be administered at any level of education, from primary to higher education, although they are more common at higher levels....
s, and then travelled in France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...
. He completed two years of national service
National service
National service is a common name for mandatory government service programmes . The term became common British usage during and for some years following the Second World War. Many young people spent one or more years in such programmes...
. 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
Ronald Fisher
Sir Ronald Aylmer Fisher FRS was an English statistician, evolutionary biologist, eugenicist and geneticist. Among other things, Fisher is well known for his contributions to statistics by creating Fisher's exact test and Fisher's equation...
's book The Genetical Theory of Natural Selection
The Genetical Theory of Natural Selection
The Genetical Theory of Natural Selection is a book by R.A. Fisher first published in 1930 by Clarendon. It is one of the most important books of the modern evolutionary synthesis and is commonly cited in biology books.-Editions:...
; but Fisher lacked standing at Cambridge as he was viewed as only a statistician. Hamilton was excited by Fisher's chapters on eugenics
Eugenics
Eugenics is the "applied science or the bio-social movement which advocates the use of practices aimed at improving the genetic composition of a population", usually referring to human populations. The origins of the concept of eugenics began with certain interpretations of Mendelian inheritance,...
. 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 his getting only a 2:1 degree.
Hamilton's rule
Hamilton enrolled in an MScMaster of Science
A Master of Science is a postgraduate academic master's degree awarded by universities in many countries. The degree is typically studied for in the sciences including the social sciences.-Brazil, Argentina and Uruguay:...
course in human demographics at the London School of Economics
London School of Economics
The London School of Economics and Political Science is a public research university specialised in the social sciences located in London, United Kingdom, and a constituent college of the federal University of London...
(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
John Hajnal
John Hajnal, FBA , born John Hajnal-Kónyi, was a Hungarian-British academic in the fields of mathematics and economics .-Life:...
of the LSE and Cedric Smith of University College London
University College London
University College London is a public research university located in London, United Kingdom and the oldest and largest constituent college of the federal University of London...
(UCL).
Both Fisher and J. B. S. Haldane
J. B. S. Haldane
John Burdon Sanderson Haldane FRS , known as Jack , was a British-born geneticist and evolutionary biologist. A staunch Marxist, he was critical of Britain's role in the Suez Crisis, and chose to leave Oxford and moved to India and became an Indian citizen...
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
Sewall Wright
Sewall Green Wright was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. With R. A. Fisher and J.B.S. Haldane, he was a founder of theoretical population genetics. He is the discoverer of the inbreeding coefficient and of...
's coefficient of relationship
Coefficient of relationship
In population genetics, Sewall Wright's coefficient of relationship or coefficient of relatedness or relatedness or r is defined as 2 times the Coefficient of Inbreeding...
. 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 fecundityFecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...
. His two 1964 papers entitled The Genetical 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
John Maynard Smith
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....
, 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
Journal of Theoretical Biology
The Journal of Theoretical Biology is a scientific journal about theoretical biology; dealing with theoretical issues, as well as mathematical and computational aspects of biology. Some research areas covered by the papers published in the journal are population genetics, morphogenesis,...
and, when first published, was largely ignored. Recognition of its significance gradually increased to the point, and it is now routinely cited in biology books. Although the theory can be proven mathematically, to date no empirical studies have calculated values for R, B, and C to determine if Hamilton's rule is ever satisfied in nature. After more than 40 years, some scientists claim that the theory remains unconfirmed, although predictions based upon it are largely supported. Because B and C represent gains and losses, it is impossible to compare actual gains and losses with the possible ones that would result from an alternate history. Statisticians, however, argue that such comparisons can be made between similar families in which different versions of history play out.
Much of the discussion relates to the evolution of eusociality
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....
in insects of the order Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...
(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 1978 Hamilton was a lecturer at University College LondonUniversity College London
University College London is a public research university located in London, United Kingdom and the oldest and largest constituent college of the federal University of London...
. Whilst there he published a paper in Science
Science (journal)
Science is the academic journal of the American Association for the Advancement of Science and is one of the world's top scientific journals....
on "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary" sex ratio
Sex ratio
Sex ratio is the ratio of males to females in a population. The primary sex ratio is the ratio at the time of conception, secondary sex ratio is the ratio at time of birth, and tertiary sex ratio is the ratio of mature organisms....
s 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
Natural history
Natural history is the scientific research of plants or animals, leaning more towards observational rather than experimental methods of study, and encompasses more research published in magazines than in academic journals. Grouped among the natural sciences, natural history is the systematic study...
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
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....
and George R. Price
George R. Price
George Robert Price was an American population geneticist. Originally a physical chemist and later a science journalist, he moved to London in 1967, where he worked in theoretical biology at the Galton Laboratory, making three important contributions: first, rederiving W.D...
were to develop into the evolutionarily stable strategy
Evolutionarily stable strategy
In game theory and behavioural ecology, an evolutionarily stable strategy , which is sometimes also called an evolutionary stable strategy, is a strategy which, if adopted by a population of players, cannot be invaded by any alternative strategy that is initially rare. An ESS is an equilibrium...
(ESS), a concept in game theory
Game theory
Game theory is a mathematical method for analyzing calculated circumstances, such as in games, where a person’s success is based upon the choices of others...
not limited to evolutionary biology. Price had originally come to Hamilton after deriving the Price equation
Price equation
The Price equation is a covariance equation which is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to re-derive W.D. Hamilton's work on kin selection...
, 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
Richard Dawkins
Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...
in Dawkins' 1976 book The Selfish Gene
The Selfish Gene
The Selfish Gene is a book on evolution by Richard Dawkins, published in 1976. It builds upon the principal theory of George C. Williams's first book Adaptation and Natural Selection. Dawkins coined the term "selfish gene" as a way of expressing the gene-centred view of evolution as opposed to the...
.
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
Harvard University
Harvard University is a private Ivy League university located in Cambridge, Massachusetts, United States, established in 1636 by the Massachusetts legislature. Harvard is the oldest institution of higher learning in the United States and the first corporation chartered in the country...
and later spent nine months with the Royal Society
Royal Society
The Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
's and the Royal Geographical Society
Royal Geographical Society
The Royal Geographical Society is a British learned society founded in 1830 for the advancement of geographical sciences...
's Xavantina-Cachimbo Expedition as a visiting professor at the University of São Paulo
University of São Paulo
Universidade de São Paulo is a public university in the Brazilian state of São Paulo. It is the largest Brazilian university and one of the country's most prestigious...
.
From 1978 Hamilton was Professor of Evolutionary Biology at the University of Michigan
University of Michigan
The University of Michigan is a public research university located in Ann Arbor, Michigan in the United States. It is the state's oldest university and the flagship campus of the University of Michigan...
. Simultaneously, he was elected a Foreign Honorary Member of American Academy of Arts and Sciences
American Academy of Arts and Sciences
The American Academy of Arts and Sciences is an independent policy research center that conducts multidisciplinary studies of complex and emerging problems. The Academy’s elected members are leaders in the academic disciplines, the arts, business, and public affairs.James Bowdoin, John Adams, and...
. His arrival sparked protests and sit-ins from students who did not like his association with sociobiology
Sociobiology
Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...
. There he worked with the political scientist
Political science
Political Science is a social science discipline concerned with the study of the state, government and politics. Aristotle defined it as the study of the state. It deals extensively with the theory and practice of politics, and the analysis of political systems and political behavior...
Robert Axelrod
Robert Axelrod
Robert M. Axelrod is an American political scientist. He is Professor of Political Science and Public Policy at the University of Michigan where he has been since 1974. He is best known for his interdisciplinary work on the evolution of cooperation, which has been cited in numerous articles...
on the prisoner's dilemma
Prisoner's dilemma
The prisoner’s dilemma is a canonical example of a game, analyzed in game theory that shows why two individuals might not cooperate, even if it appears that it is in their best interest to do so. It was originally framed by Merrill Flood and Melvin Dresher working at RAND in 1950. Albert W...
,
and was a member of the BACH group with original members Arthur Burks
Arthur Burks
Arthur Walter Burks was an American mathematician who in the 1940s as a senior engineer on the project contributed to the design of the ENIAC, the first general-purpose electronic digital computer. Decades later, Burks and his wife Alice Burks outlined their case for the subject matter of the...
, Robert Axelrod
Robert Axelrod
Robert M. Axelrod is an American political scientist. He is Professor of Political Science and Public Policy at the University of Michigan where he has been since 1974. He is best known for his interdisciplinary work on the evolution of cooperation, which has been cited in numerous articles...
, Michael Cohen and John Holland
John Henry Holland
John Henry Holland is an American scientist and Professor of Psychology and Professor of Electrical Engineering and Computer Science at the University of Michigan, Ann Arbor. He is a pioneer in complex systems and nonlinear science. He is known as the father of genetic algorithms. He was awarded...
.
Chasing the Red Queen
Hamilton was an early proponent of the Red QueenRed Queen's Hypothesis
The Red Queen's Hypothesis, also referred to as Red Queen, Red Queen's race or Red Queen Effect, is an evolutionary hypothesis. The term is taken from the Red Queen's race in Lewis Carroll's Through the Looking-Glass...
theory of the evolution of sex
Evolution of sex
The evolution of sexual reproduction is currently described by several competing scientific hypotheses. All sexually reproducing organisms derive from a common ancestor which was a single celled eukaryotic species. Many protists reproduce sexually, as do the multicellular plants, animals, and fungi...
, first proposed by Leigh Van Valen
Leigh Van Valen
Leigh Maiorana Van Valen was an American evolutionary biologist. He was professor emeritus in the Department of Ecology and Evolution at the University of Chicago....
. This was named for a character in Lewis Carroll
Lewis Carroll
Charles Lutwidge Dodgson , better known by the pseudonym Lewis Carroll , was an English author, mathematician, logician, Anglican deacon and photographer. His most famous writings are Alice's Adventures in Wonderland and its sequel Through the Looking-Glass, as well as the poems "The Hunting of the...
's Through the Looking-Glass
Through the Looking-Glass
Through the Looking-Glass, and What Alice Found There is a work of literature by Lewis Carroll . It is the sequel to Alice's Adventures in Wonderland...
, 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
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
, 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 SouthwoodRichard Southwood
Sir Thomas Richard Edmund Southwood DL, FRS was Professor of zoology and Vice-Chancellor of the University of Oxford....
to be the Royal Society Research Professor in the Department of Zoology at Oxford, and a Fellow of New College
New College, Oxford
New College is one of the constituent colleges of the University of Oxford in the United Kingdom.- Overview :The College's official name, College of St Mary, is the same as that of the older Oriel College; hence, it has been referred to as the "New College of St Mary", and is now almost always...
, where he remained until his death.
From 1994 Hamilton found companionship with Maria Luisa Bozzi, an Italian
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...
science journalist
Journalist
A journalist collects and distributes news and other information. A journalist's work is referred to as journalism.A reporter is a type of journalist who researchs, writes, and reports on information to be presented in mass media, including print media , electronic media , and digital media A...
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 evolutionSocial evolution
Social evolution is a subdiscipline of evolutionary biology that is concerned with social behaviors that have fitness consequences for individuals other than the actor...
, 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.
Through his collaboration with Hugh N. Comins and Bob May
Robert May, Baron May of Oxford
Robert McCredie May, Baron May of Oxford, OM, AC, PRS is an Australian scientist who has been Chief Scientific Adviser to the UK Government, President of the Royal Society, and a Professor at Sydney and Princeton. He now holds joint professorships at Oxford, and Imperial College London...
on evolutionarily stable dispersal strategies, Hamilton acquired an Erdős number
Erdos number
The Erdős number describes the "collaborative distance" between a person and mathematician Paul Erdős, as measured by authorship of mathematical papers.The same principle has been proposed for other eminent persons in other fields.- Overview :...
of 5.
Hamilton also proposed the coevolution theory of autumn leaf color
Autumn leaf color
Autumn leaf color is a phenomenon that affects the normally green leaves of many deciduous trees and shrubs by which they take on, during a few weeks in the autumn season, one or many colors that range from red to yellow...
as an example of evolutionary signalling theory
Signalling theory
Within evolutionary biology, signalling theory is a body of theoretical work examining communication between individuals. The central question is when organisms with conflicting interests should be expected to communicate "honestly"...
.
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 KoprowskiHilary Koprowski
Hilary Koprowski is a Polish virologist and immunologist, and inventor of the world's first effective live polio vaccine.-Life:...
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
OPV AIDS hypothesis
The oral polio vaccine AIDS hypothesis argues that the AIDS pandemic originated from live polio vaccines prepared in chimpanzee tissue cultures and then administered to up to one million Africans between 1957 and 1960 in experimental mass vaccination campaigns.Data from molecular biology and...
by assessing natural levels of SIV in primates, in early 2000 he and two others ventured on a field trip to the war-torn Democratic Republic of the Congo
Democratic Republic of the Congo
The Democratic Republic of the Congo is a state located in Central Africa. It is the second largest country in Africa by area and the eleventh largest in the world...
.
Death
He returned to London from Africa on 29 Jan 2000. He was admitted to University College Hospital London on 30 Jan 2000. He was transferred to Middlesex Hospital London on 5 Feb 2000 and died there on 7 Mar 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".A secular memorial service (he was an atheist) was held at the Chapel of New College, Oxford
New College, Oxford
New College is one of the constituent colleges of the University of Oxford in the United Kingdom.- Overview :The College's official name, College of St Mary, is the same as that of the older Oriel College; hence, it has been referred to as the "New College of St Mary", and is now almost always...
on Saturday 1 July 2000, organised by Richard Dawkins
Richard Dawkins
Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...
. He was buried near Wytham Woods
Wytham
Wytham is a village and civil parish on Seacourt Stream, a branch of the River Thames, about northwest of Oxford. It is just west of the Western By-Pass Road, part of the Oxford Ring Road ....
. He, however, had written an essay on My intended burial and why in which he wrote:
The second volume of his collected papers, Evolution of Sex, was published in 2002, and the third and final volume, Last Words, in 2005.
Awards
- 1978 Foreign Honorary Member of American Academy of Arts and SciencesAmerican Academy of Arts and SciencesThe American Academy of Arts and Sciences is an independent policy research center that conducts multidisciplinary studies of complex and emerging problems. The Academy’s elected members are leaders in the academic disciplines, the arts, business, and public affairs.James Bowdoin, John Adams, and...
- 1980 Fellow of the Royal SocietyRoyal SocietyThe Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
of London - 1982 Newcomb Cleveland PrizeNewcomb Cleveland PrizeThe Newcomb Cleveland Prize of the American Association for the Advancement of Science is annually awarded to author of outstanding scientific paper published in the Research Articles or Reports sections of Science...
of the American Association for the Advancement of ScienceAmerican Association for the Advancement of ScienceThe American Association for the Advancement of Science is an international non-profit organization with the stated goals of promoting cooperation among scientists, defending scientific freedom, encouraging scientific responsibility, and supporting scientific education and science outreach for the... - 1988 Darwin MedalDarwin MedalThe Darwin Medal is awarded by the Royal Society every alternate year for "work of acknowledged distinction in the broad area of biology in which Charles Darwin worked, notably in evolution, population biology, organismal biology and biological diversity". First awarded in 1890, it was created in...
of the Royal SocietyRoyal SocietyThe Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
of London - 1989 Scientific Medal of the Linnean Society
- 1991 Frink MedalFrink MedalThe Frink Medal for British Zoologists is awarded by the Zoological Society of London "For significant and original contributions by a professional zoologist to the development of zoology in the wider applications."- Recipients :...
of Zoological Society of London - 1992/3 Wander Prize of the University of Bern
- 1993 Crafoord PrizeCrafoord PrizeThe Crafoord Prize is an annual science prize established in 1980 by Holger Crafoord, a Swedish industrialist, and his wife Anna-Greta Crafoord...
of the Royal Swedish Academy of SciencesRoyal Swedish Academy of SciencesThe Royal Swedish Academy of Sciences or Kungliga Vetenskapsakademien is one of the Royal Academies of Sweden. The Academy is an independent, non-governmental scientific organization which acts to promote the sciences, primarily the natural sciences and mathematics.The Academy was founded on 2... - 1993 Kyoto PrizeKyoto PrizeThe has been awarded annually since 1985 by the Inamori Foundation, founded by Kazuo Inamori. The prize is a Japanese award similar in intent to the Nobel Prize, as it recognizes outstanding works in the fields of philosophy, arts, science and technology...
of the Inamori Foundation - 1995 Fyssen Prize of the Fyssen FoundationFyssen FoundationThe Fyssen Foundation is a French charitable organization that was established and endowed in 1979 by H. Fyssen. The aim of the foundation is to stimulate research into the processes underlying and leading to cognition, including work in such disciplines as ethology, paleontology, archaeology,...
Biographies
- Alan GrafenAlan GrafenAlan Grafen is a Scottish ethologist and evolutionary biologist. He currently teaches and undertakes research at St John's College, Oxford. Along with regular contributions to scientific journals, Grafen is known publicly for his work as co-editor of the 2006 festschrift Richard Dawkins: How a...
has written a biographical memoir for the Royal Society. - A book is also in press: Segerstråle, 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
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
- 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. with Robert AxelrodRobert AxelrodRobert M. Axelrod is an American political scientist. He is Professor of Political Science and Public Policy at the University of Michigan where he has been since 1974. He is best known for his interdisciplinary work on the evolution of cooperation, which has been cited in numerous articles...
Other references
- Edwards A.W.F.A. W. F. EdwardsAnthony William Fairbank Edwards is a British statistician, geneticist, and evolutionary biologist, sometimes called Fisher's Edwards. He is a Life Fellow of Gonville and Caius College and retired Professor of Biometry at the University of Cambridge, and holds both the ScD and LittD degrees. A...
(1998) Notes and Comments. - Fisher R.A.Ronald FisherSir Ronald Aylmer Fisher FRS was an English statistician, evolutionary biologist, eugenicist and geneticist. Among other things, Fisher is well known for his contributions to statistics by creating Fisher's exact test and Fisher's equation...
(1930). The Genetical Theory of Natural SelectionThe Genetical Theory of Natural SelectionThe Genetical Theory of Natural Selection is a book by R.A. Fisher first published in 1930 by Clarendon. It is one of the most important books of the modern evolutionary synthesis and is commonly cited in biology books.-Editions:...
. Clarendon Press, Oxford. - Ford E.B.E.B. FordEdmund Brisco "Henry" Ford FRS Hon. FRCP was a British ecological geneticist. He was a leader among those British biologists who investigated the role of natural selection in nature. As a schoolboy Ford became interested in lepidoptera, the group of insects which includes butterflies and moths...
(1945) New Naturalist 1: Butterflies. Collins: London. - Maynard Smith J.John Maynard SmithJohn Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....
and G.R. PriceGeorge R. PriceGeorge Robert Price was an American population geneticist. Originally a physical chemist and later a science journalist, he moved to London in 1967, where he worked in theoretical biology at the Galton Laboratory, making three important contributions: first, rederiving W.D...
(1973) The logic of animal conflict. NatureNature (journal)Nature, first published on 4 November 1869, is ranked the world's most cited interdisciplinary scientific journal by the Science Edition of the 2010 Journal Citation Reports...
146: 15—18. - Dawkins R.Richard DawkinsClinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...
(1989) The Selfish GeneThe Selfish GeneThe Selfish Gene is a book on evolution by Richard Dawkins, published in 1976. It builds upon the principal theory of George C. Williams's first book Adaptation and Natural Selection. Dawkins coined the term "selfish gene" as a way of expressing the gene-centred view of evolution as opposed to the...
, 2nd ed. Oxford University Press. - Madsen E.A., Tunney R. Fieldman, G. Plotkin H.C., Dunbar R.I.M.Robin DunbarRobin Ian MacDonald Dunbar is a British anthropologist and evolutionary psychologist and a specialist in primate behaviour. He is currently Professor of Evolutionary Anthropology and the Director of the Institute of Cognitive and Evolutionary Anthropology of the University of Oxford and the...
, Richardson J.M. & McFarland D. (2006) Kinship and altruism: a cross-cultural experimental study. British Journal of Psychology
Other references
- 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 Hamilton
Game Theory