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For other persons named William Hamilton, see William Hamilton (disambiguation). William Hamilton (and shortened forms may refer to Europeans William Hamilton (Lord Chancellor, (d
W. D. Hamilton, 1996
W. D. Hamilton, 1996

William Donald Hamilton, F.R.S. (1 August 1936 — 7 March 2000) was a British evolutionary biologist, considered one of the greatest evolutionary theorists of the 20th century. The Royal Society of London for the Improvement of Natural Knowledge, known simply as The Royal Society, is a Learned society for science that was founded in 1660 Events 30 BC - Octavian (later known as Augustus enters Alexandria, Egypt, bringing it under the control of the Roman Year 1936 ( MCMXXXVI) was a Leap year starting on Wednesday (link will display the full calendar of the Gregorian calendar. Events 161 - Roman Emperor Antoninus Pius dies and is succeeded by co-Emperors Marcus Aurelius and Lucius Verus 2000 ( MM) was a Leap year that started on Saturday of the Common Era, in accordance with the Gregorian calendar. The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom, the UK or Britain,is a Sovereign state located Evolutionary biology is a sub-field of Biology concerned with the origin of Species from a Common descent, and Descent of species eVolution is the third Album by eLDee, it was due to be released in 2008 [1] From 1984 to his death in 2000, he was the Royal Society Research Professor at Oxford University. The Royal Society of London for the Improvement of Natural Knowledge, known simply as The Royal Society, is a Learned society for science that was founded in 1660 The University of Oxford (informally "Oxford University" or simply "Oxford" located in the city of Oxford, Oxfordshire, England is the Hamilton became famous for his theoretical work expounding a rigorous genetic basis for the existence of kin selection. Theoretical biology is a field of academic study and research that involves the use of Models and theories in Biology. Genetics (from Ancient Greek grc-Latn genetikos, “genitive” and that from grc-Latn genesis, “origin” a discipline of Biology, is From the time of antiquity field biologists have observed that some organisms tend to exhibit strategies that favor the reproductive success of their relatives even at a cost to their own survival This insight was a key part of the development of a gene-centric view of evolution, and he can therefore be seen as one of the forerunners of the discipline of sociobiology popularized by E. O. Wilson. The gene-centered view of evolution, gene selection theory or selfish gene theory holds that Natural selection acts through differential survival of competing Sociobiology is a neo-Darwinian and Socialism Synthesis of Scientific disciplines that attempts to explain Social behavior Edward Osborne Wilson (born June 10, 1929) is an American biologist researcher ( Sociobiology, Biodiversity) theorist ( Hamilton also published important work on sex ratios and the evolution of sex. 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

Contents

Biography

Early life

Hamilton was born in 1936 in Cairo, Egypt, the second eldest of six children. Cairo () which means "the Vanquisher" or "the Triumphant" is the capital and largest city of Egypt. This article is about the country of Egypt For a topic outline on this subject see List of basic Egypt topics. His father A. M. Hamilton was a New Zealand-born engineer. Archibald Milne Hamilton (1898 - 1972 was a New Zealand -born Civil engineer, notable for building the Hamilton Road through Kurdistan and designing New Zealand is an Island country in the south-western Pacific Ocean comprising two main landmasses (the North Island and the South Island His mother B. M. Hamilton was a medical doctor. A physician, medical practitioner or medical doctor who practices Medicine, and is concerned with maintaining or restoring human Health

The Hamilton family settled in Kent. KENT (1400 AM) is a Radio station broadcasting a Adult Standards/MOR format During the Second World War he was evacuated to Edinburgh. World War II, or the Second World War, (often abbreviated WWII) was a global military conflict which involved a majority of the world's nations, including Edinburgh ( ˈɛdɪnb(ərə Dùn Èideann) is the Capital of Scotland and is its second largest city after Glasgow. He had an interest in natural history from an early age and would spend his spare time collecting butterflies and other insects. A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a 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. Professor Edmund Brisco "Henry" Ford FRS, Hon FRCP ( 23rd April 1901- 2nd January 1988 was a British ecological The New Naturalist books are a series published by Collins in the United Kingdom, on a variety of Natural history topics relevant to the British Isles Natural selection is the process by which favorable Heritable traits become more common in successive Generations of a Population of Genetics (from Ancient Greek grc-Latn genetikos, “genitive” and that from grc-Latn genesis, “origin” a discipline of Biology, is Population genetics is the study of the Allele frequency distribution and change under the influence of the four evolutionary forces Natural selection, Genetic

He was educated at Tonbridge School, where he was in the School House. Tonbridge School is a British independent all boys Public school in Tonbridge, founded in 1553 by Sir Andrew Judde As a 12-year old he was seriously injured while playing with explosives his father had left over from when he made hand grenades for the Home Guard during the Second World War, an accident that probably would have killed him had his mother not been medically qualified. An explosive material is a material that either is chemically or otherwise Energetically unstable or produces a sudden expansion of the material usually accompanied The British Home Guard (initially "Local Defence Volunteers" or LDV, or in slang Look-Duck-Vanish, hence the name change was a defence World War II, or the Second World War, (often abbreviated WWII) was a global military conflict which involved a majority of the world's nations, including A thoracotomy in King’s College Hospital saved his life, but the explosion left him with amputated fingers on his right hand and scarring on his body — he took six months to recover. Thoracotomy is an incision into the Chest. It is performed by a surgeon and rarely by emergency physicians and Paramedics, to gain access to the thoracic organs

Hamilton stayed on an extra term at Tonbridge in order to complete the Cambridge entrance examinations, and then travelled in France. The University of Cambridge (often Cambridge University) located in Cambridge, England, is the second-oldest university in the An entrance examination is an Examination that many educational institutions use to select students for admission This article is about the country For a topic outline on this subject see List of basic France topics. He then completed two years of national service. National service is a common name for mandatory or voluntary government service programs (most often focusing on military service As an undergraduate at St. John's College, he was uninspired by the fact that there "many biologists hardly seemed to believe in evolution". In some Educational systems undergraduate education is Post-secondary education up to the level of a Bachelor's degree. St John's College, an institution known formally as The Master Fellows and Scholars of the College of St John the Evangelist in the University of Cambridge is a Nevertheless, he came across Ronald Fisher's book The Genetical Theory of Natural Selection; Fisher lacked standing at Cambridge as he was viewed only as a statistician. Sir Ronald Aylmer Fisher, FRS ( 17 February 1890 – 29 July 1962) was an English Statistician, Evolutionary The Genetical Theory of Natural Selection is a book by RA Fisher. Hamilton wrote on a postcard to his sister Mary on the day he found the book, excited by its chapters on eugenics. Eugenics is a social Philosophy which advocates the improvement of Human Hereditary traits through various forms of intervention 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

Main article: Hamilton's rule

Hamilton having various ideas and problems enrolled on an MSc course in human demographics at the London School of Economics (LSE), under Norman Carrier who secured for him various grants. The British undergraduate degree classification system is a grading scheme for Undergraduate degrees ( Bachelor's degrees and some Master's degrees From the time of antiquity field biologists have observed that some organisms tend to exhibit strategies that favor the reproductive success of their relatives even at a cost to their own survival The London School of Economics and Political Science, more commonly referred to as The London School of Economics or LSE, is a specialist college of the Later when work became more mathematical and then genetical, he had his supervision transferred to John Hajnal of the LSE and Cedric Smith of University College London (UCL). John Hajnal (born 26 November 1924) was Professor of Statistics London School of Economics, 1975-86 Cedric Austen Bardell Smith ( February 5 1917 &ndash January 10 2002) was a British Statistician and Geneticist University College London ( UCL) is a multi-faculty university institution based in the United Kingdom and a constituent college of the University of London

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. John Burdon Sanderson Haldane FRS ( 5 November 1892 &ndash 1 December 1964) known as Jack (but who used 'J 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. Sewall Green Wright ( December 21, 1889 – March 3, 1988) was an American Geneticist In Population genetics, Sewall Wright 's coefficient of relationship or coefficient of relatedness or relatedness or r is Thus became Hamilton's rule. Briefly, the rule is that a costly action should be performed if:

C < R \times 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. His two 1964 papers entitled The Genetical Evolution of Social Behavior are now widely referenced. The Genetical Evolution of Social Behavior is a 1964 Scientific paper by the British Evolutionary biologist W

The proof and discussion of its consequences however involved heavy mathematics, and was passed over by two reviewers. The third, John Maynard Smith, did not completely understand it either, but recognised its significance; this passing over would later lead to friction between Hamilton and Maynard Smith, Hamilton feeling that Maynard Smith had held his work back to claim credit for the idea himself. Maynard Smith redirects here -- for other uses see Maynard Smith (disambiguation Professor John Maynard Smith, F The paper was printed in the relatively obscure Journal of Theoretical Biology, and when first published was largely ignored. The Journal of Theoretical Biology is a Scientific journal about Theoretical biology; dealing with theoretical issues as well as mathematical The significance of it gradually increased, to the point where they are routinely cited in biology books. To date, however, there have been no empirical studies that have calculated values for R, B, and C to determine if Hamilton's rule is ever satisfied in nature; as such, even after more than 40 years, the theory remains unproven, though predictions based upon the theory are largely supported.

A large part of the discussion related to the evolution of eusociality in insects of the order Hymenoptera (ants, bees and wasps) based on their unusual haplodiploid sex-determination system. Eusociality ( Greek eu: "good" + "social" is a term used for the highest level of social organization in a hierarchical classification Hymenoptera is one of the larger orders of Insects comprising the sawflies, Wasps Bees and Ants The name refers to Ants are social Insects of the family Formicidae and along with the related families of Wasps and Bees belong to the order Bees are flying Insects closely related to Wasps and Ants Bees are a Monophyletic lineage within the superfamily Apoidea A wasp is any Insect of the order Hymenoptera and suborder Apocrita that is neither a Bee nor Ant. The haplodiploid sex-determination system determines the sex of the offspring of many Hymenopterans ( Bees Ants and Wasps, and coleopterans ( This system means that females are more closely related to their sisters than to their own (potential) offspring. Thus, Hamilton reasoned, a "costly action" would be better spent in helping to raise their sisters, rather than reproducing themselves.

Extraordinary sex ratios

Between 1964 and 1978 Hamilton was a lecturer at Imperial College London. Imperial College London (officially The Imperial College of Science Technology and Medicine as given in its Royal Charter It is one of only three universities to have reached Whilst there he published a paper in Science on "extraordinary sex ratios". Science is the Academic journal of the American Association for the Advancement of Science and is considered one of the world's most prestigious Scientific 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. 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 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. Anthony William Fairbank Edwards (born 1935 is a British Statistician, Geneticist, and Evolutionary biologist. Hamilton combined his extensive knowledge of natural history with deep insight into the problem, opening up a whole new area of research. Natural history is the Scientific research of Plants or Animals leaning more towards the Observational than Experimental methods

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. Maynard Smith redirects here -- for other uses see Maynard Smith (disambiguation Professor John Maynard Smith, F George R Price (1922 – January 6, 1975) was an American population geneticist. In Game theory and Behavioural ecology, an evolutionarily stable strategy (ESS is a strategy which if adopted by a population of players Game theory is a branch of Applied mathematics that is used in the Social sciences (most notably Economics) Biology, Engineering, Price had originally come to Hamilton after deriving the Price equation, and thus rederiving Hamilton's rule. The Price equation (also known as Price's equation) is a covariance equation which is a mathematical description of Evolution and Natural selection. 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. Clinton Richard Dawkins, FRS, FRSL (born 26 March 1941 is a British ethologist, evolutionary biologist, and Popular science

In 1966 he married Christine Friess and they were to have three daughters, Helen, Ruth and Rowena. 26 years later they amicably separated.

Hamilton was a visiting professor at Harvard University and later spent nine months with the Royal Society's and the Royal Geographic Society's Xavantina-Cachimbo Expedition as a visiting professor at the University of São Paulo. The Royal Society of London for the Improvement of Natural Knowledge, known simply as The Royal Society, is a Learned society for science that was founded in 1660 History Founding members of the Society include Sir John Barrow, Sir John Franklin and Francis Beaufort. The University of São Paulo (in Portuguese Universidade de São Paulo; USP) is one of the three public universities funded by the State of São Paulo.

From 1978 Hamilton was Professor of Evolutionary Biology at the University of Michigan. Evolutionary biology is a sub-field of Biology concerned with the origin of Species from a Common descent, and Descent of species The University of Michigan Ann Arbor ( U of M, U-M, UM or simply Michigan) is a top-ranked Coeducational public research Simultaneously, he was elected a Foreign Honorary Member of American Academy of Arts and Sciences. The American Academy of Arts and Sciences (AAAS is an organization dedicated to scholarship and the advancement of learning His arrival sparked protests and sit-ins from students who did not like his association with sociobiology. A sit-in or sit-down is a form of direct action that involves one or more persons nonviolently occupying an area for a Protest, often to promote political social Sociobiology is a neo-Darwinian and Socialism Synthesis of Scientific disciplines that attempts to explain Social behavior 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. This is a list of notable political scientists See the List of political theorists for those who study politics without using the Scientific method. Robert Axelrod (born 1943 is a Professor of Political Science and Public Policy at the University of Michigan. The Prisoner's Dilemma constitutes a problem in Game theory. It was originally framed by Merrill Flood and Melvin Dresher Arthur Walter Burks ( October 13 1915 – May 14, 2008) was an American Mathematician who in the 1940s as a senior Robert Axelrod (born 1943 is a Professor of Political Science and Public Policy at the University of Michigan. John Henry Holland ( 2 February, 1929) is an American scientist and Professor of Psychology and Professor of Electrical Engineering and Computer

Chasing the Red Queen

Hamilton was an early proponent of the Red Queen theory of the evolution of sex,[2] first proposed by Leigh Van Valen. The Red Queen's Hypothesis, Red Queen, " Red Queen's race " or " Red Queen Effect " is an Evolutionary Hypothesis Professor Leigh M Van Valen (born 1935) is an American evolutionary biologist. 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. Charles Lutwidge Dodgson (ˈdɒdsən (27 January 1832 &ndash 14 January 1898 better known by the Pen name Lewis Carroll (/ˈkærəl/ was an English Through the Looking-Glass and What Alice Found There ( 1871) is a work of Children's literature by Lewis Carroll (Charles Lutwidge Dodgson "
"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. Parasitism is a type of symbiotic relationship between Organisms of different Species. Likewise, parasites were able to evolve mechanisms to get around the organism's new set of genes, thus perpetuating an endless race.

Back in Britain

In 1980 he was elected a Fellow of the Royal Society, and in 1984 he was invited by Richard Southwood to be the Royal Society Research Professor at New College, Oxford, Department of Zoology, where he remained until his death. The Royal Society of London for the Improvement of Natural Knowledge, known simply as The Royal Society, is a Learned society for science that was founded in 1660 Sir Thomas Richard Edmund Southwood DL, FRS ( 20 June 1931 &ndash 26 October 2005) was Professor of Zoology New College is one of the constituent colleges of the University of Oxford in the United Kingdom. The University of Oxford (informally "Oxford University" or simply "Oxford" located in the city of Oxford, Oxfordshire, England is the

From 1994 Hamilton found companionship with Maria Luisa Bozzi, an Italian science journalist and author. Italy (Italia officially the Italian Republic, (Repubblica Italiana is located on the Italian Peninsula in Southern Europe, and on the two largest A journalist (also called a newspaperman) is a person who practices Journalism, the gathering and dissemination of information about current events trends

On the Origin of HIV

During the 1990s Hamilton became increasingly convinced by the controversial argument that the origin of the HIV virus lay in oral polio vaccines (the OPV AIDS hypothesis) in Africa during the 1950s. Human immunodeficiency virus ( HIV) is a Lentivirus (a member of the Retrovirus family that can lead to acquired immunodeficiency syndrome Two polio Vaccines are used throughout the world to combat Poliomyelitis (or polio See also AIDS origin The oral polio vaccine (OPV AIDS hypothesis argues that the AIDS Pandemic originated from live Polio vaccines prepared Letters by Hamilton to Science were rejected by the journal, amid accusations that the medical establishment were ranging against the OPV hypothesis. Science is the Academic journal of the American Association for the Advancement of Science and is considered one of the world's most prestigious Scientific

To find indirect evidence of the OPV hypothesis by assessing natural levels of SIV in primates, he and two others ventured on a field trip to the war-torn Democratic Republic of the Congo, where he contracted malaria. The Democratic Republic of the Congo (République démocratique du Congo often referred to as DR Congo, DRC or RDC, and formerly known or referred to Malaria is a vector -borne Infectious disease caused by Protozoan Parasites It is widespread in tropical and subtropical regions including He was rushed home and spent six weeks in hospital before dying from a cerebral haemorrhage.

Postscript

A secular memorial service (he was an atheist) was held at the Chapel of New College, Oxford on Saturday 1 July 2000, organised by Richard Dawkins. Atheism New College is one of the constituent colleges of the University of Oxford in the United Kingdom. "July 1st" redirects here For the Ayumi Hamasaki song see H (song. 2000 ( MM) was a Leap year that started on Saturday of the Common Era, in accordance with the Gregorian calendar. Clinton Richard Dawkins, FRS, FRSL (born 26 March 1941 is a British ethologist, evolutionary biologist, and Popular science

His body was interred in Wytham Woods. Wytham (ˈwaɪtəm is a small village in central Oxfordshire on the south bank of the Seacourt Stream, a branch of the River Thames, three miles west from He however had written an essay on My intended burial and why in which he wrote:[3]

“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 was published in 2002.

Social evolution

The field of social evolution, of which Hamilton's rule has central importance, is broadly defined as being the study of the evolution of social behaviours, i. Social Evolution is the title of an essay by Benjamin Kidd, which became available as a book published by Macmillan and co London in 1894 From the time of antiquity field biologists have observed that some organisms tend to exhibit strategies that favor the reproductive success of their relatives even at a cost to their own survival 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. Robert McCredie May Baron May of Oxford, OM, AC, FRS (born Australia, 8 January 1936) has been Chief Scientific Adviser The Erdős number (ɛrdøːʃ honoring the late Hungarian mathematician Paul Erdős, is a way of describing the "collaborative distance" between a person

Awards

Biographies

Works

Collected papers

Hamilton started to publish his collected papers starting in 1996, along the lines of Fisher's collected papers, with short essays giving each paper context. He died after the preparation of the second volume, so the essays for the third volume come from his coauthors.

Significant papers

Other references

Cited references

  1. ^ Obituary by Richard Dawkins - The Independent - 10 March 2000
  2. ^ The Red Queen Hypothesis at Indiana University. Quote: "W. D. Hamilton and John Jaenike were among the earliest pioneers of the idea. "
  3. ^ Hamilton, W. D. (2000) My intended burial and why, Ethology Ecology and Evolution 12 111-122 PDF

Other references

Game Theory

AIDS

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