W. D. Hamilton

William Donald "Bill" Hamilton, F.R.S. (1 August 1936 — 7 March 2000) was a British evolutionary biologist, considered one of the greatest evolutionary theorists of the 20th century.[1] Hamilton became famous for his theoretical work expounding a rigorous genetic basis for the existence of kin selection. This insight was a key part of the development of a gene-centric view of evolution, and he can therefore be seen as one of the forerunners of the discipline of sociobiology founded by Edward Osborne Wilson. Hamilton also published important work on sex ratios and the evolution of sex. Image File history File links W_D_Hamilton2. ... The premises of the Royal Society in London (first four properties only). ... August 1 is the 213th day of the year in the Gregorian Calendar (214th in leap years), with 152 days remaining. ... 1936 (MCMXXXVI) was a leap year starting on Wednesday (link will take you to calendar). ... March 7 is the 66th day of the year in the Gregorian Calendar (67th in leap years). ... This article is about the year 2000. ... Evolutionary biology is a subfield of biology concerned with the origin and descent of species, as well as their change over time, i. ... Kin selection refers to changes in gene frequency across generations that are driven at least in part by interactions between related individuals, and this forms much of the conceptual basis of the theory of social evolution. ... This article needs to be cleaned up to conform to a higher standard of quality. ... Sociobiology is a synthesis of scientific disciplines that attempts to explain behaviour in all species by considering the evolutionary advantages of social behaviours. ... E.O. Wilson with Dynastes hercules E. O. Wilson, or Edward Osborne Wilson, (born June 10, 1929) is an entomologist and biologist known for his work on ecology, evolution, and sociobiology. ... Sex ratio by country for total population Sex ratio is the ratio of males to females in a population. ... The evolution of sex is a major puzzle in modern evolutionary biology. ...

Biography

Early life

Hamilton was born in 1936 in Cairo, Egypt, the second eldest of six children. His father, A. M. Hamilton was a New Zealand-born engineer, and his mother, B. M. Hamilton was a medical doctor. Cairo Minarets Cairo (Arabic: ‎ transliterated: , transl. ... Archibald Milne Hamilton (? - 1972) was a New Zealand-born engineer, notable for building the Hamilton Road through Kurdistan and designing the Hamilton-Callender bridge system. ... The word physician should not be confused with physicist, which means a scientist in the area of physics. ...

The Hamilton family moved to Kent when Bill was young and during the Second World War he was evacuated to Edinburgh. He had an interest in natural history from an early age and would spend his spare time collecting butterflies and other insects. In 1946 he discovered E.B. Ford's New Naturalist book Butterflies, which introduced him to the principles of evolution by natural selection, genetics and population genetics. Mushroom cloud from the nuclear explosion over Nagasaki rising 18 km into the air. ... Edinburgh (pronounced ; Dùn Èideann () in Scottish Gaelic) is Scotlands capital, and its second-largest city. ... For other uses of the term butterfly, see butterfly (disambiguation). ... Professor Edmund Brisco Henry Ford, F.R.S. (23rd April 1901 - 2nd January 1988) was a British ecological geneticist. ... Cover of the first book in the series, E.B. Fords famous Butterflies 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. ... A hypothetical phylogenetic tree of all extant organisms, based on 16S rRNA gene sequence data, showing the evolutionary history of the three domains of life, bacteria, archaea and eukaryotes. ... Natural selection is the process by which individual organisms with favorable traits are more likely to survive and reproduce. ... Genetics (from the Greek genno γεννώ= give birth) is the science of genes, heredity, and the variation of organisms. ... Population genetics is the study of the distribution of and change in allele frequencies under the influence of the four evolutionary forces: natural selection, genetic drift, mutation, and migration. ...

He was educated at Tonbridge School, where he was in the School House. As a 12-year old he was seriously injured while playing with explosives his father had left over from when he made hand grenades for the Home Guard during the Second World War, an accident that probably would have killed him had his mother not been medically qualified. A thoracotomy in King’s College Hospital saved his life, but the explosion left him with amputated fingers on his right hand and scarring on his body — he took six months to recover. Tonbridge School is a British independent all boys boarding school in Tonbridge, founded in 1553 by Sir Andrew Judde. ... This article is concerned solely with chemical explosives. ... A WWII-era MkIIA1 pineapple fragmentation hand grenade A hand grenade is a small hand-held bomb designed to be thrown by hand. ... A Home Guard is a part-time civilian reserve military force similar to a militia. ... Mushroom cloud from the nuclear explosion over Nagasaki rising 18 km into the air. ... Thoracotomy is a surgical incision into the chest. ...

Hamilton stayed on an extra term at Tonbridge in order to complete the Cambridge entrance examinations, and then travelled in France. He then completed two years of national service. As an undergraduate at St. John's College, he was uninspired by the fact that there "many biologists hardly seemed to believe in evolution". Nevertheless, he came across Ronald Fisher's book The Genetical Theory of Natural Selection; Fisher lacked standing at Cambridge as was viewed only as a statistician. Hamilton wrote on a postcard to his sister Mary on the day he found the book excited by its darker 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 only getting 2:1 degree. The University of Cambridge (often called Cambridge University, or just Cambridge), located in Cambridge, England, is the second-oldest university in the English-speaking world. ... An entrance examination is an examination that educational institutions use to determine whether prospective students are good enough to enter their institution. ... // National Service National Service in the 20th Century referred primarily to conscription for military service. ... In some educational systems, an undergraduate is a post-secondary student pursuing a Bachelors degree. ... Full name The College of Saint John the Evangelist of the University of Cambridge Motto - Named after The Hospital of Saint John the Evangelist, Cambridge, named after John the Evangelist Previous names - Established 1511 Sister College Balliol College Master Prof. ... Sir Ronald Fisher Sir Ronald Aylmer Fisher, FRS (17 February 1890 – 29 July 1962) was a British eugenicist, evolutionary biologist, geneticist and statistician. ... The Genetical Theory of Natural Selection is a book by Ronald Fisher. ... Eugenics is the self-direction of human evolution: Logo from the Second International Congress of Eugenics, 1921, depicting it as a tree which unites a variety of different fields. ... A hypothetical phylogenetic tree of all extant organisms, based on 16S rRNA gene sequence data, showing the evolutionary history of the three domains of life, bacteria, archaea and eukaryotes. ...

Hamilton's rule

Hamilton having various ideas and problems enrolled on an MSc course in human demographics at the London School of Economics (LSE), under Norman Carrier who secured for him various grants. Later when work became more mathematical and then genetical, he had his supervision transferred to John Hajnal of the LSE and Cedric Smith of University College London (UCL). The London School of Economics and Political Science, often referred to as the London School of Economics or simply the LSE, is a specialist university and a constituent college of the federal University of London, located on Houghton Street in Central London, off the Aldwych and next to the Royal... Cedric Austen Bardell Smith (February 5, 1917, — January 16, 2002) was a British statistician Categories: Scientist stubs | 1917 births | 2002 deaths ... University College London, commonly known as UCL, is one of the colleges that make up 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. Hamilton worked through several examples, and eventually realised that the number that kept falling out of his calculations was Sewall Wright's coefficient of relationship. Thus became Hamilton's rule. Briefly, the rule is that a costly action should be performed if; John Burdon Sanderson Haldane (November 5, 1892 - December 1, 1964), who normally used J.B.S. as a first name, was a geneticist born in Scotland and educated at Eton and Oxford University. ... Sewall Green Wright ForMemRS (December 21, 1889 – March 3, 1988) was an American geneticist known for his influential work on evolutionary theory. ... In population genetics, Sewall Wrights coefficient of relationship or relatedness is the probability that at a random locus, the alleles there will be identical by descent. ...

Where C is the cost to the actor, R the genetic relatedness between the actor and the recipient and B is the benefit to the recipient. Costs and benefits are measured in fecundity. His two 1964 papers entitled The Evolution of Social Behaviour are now universally referenced. The Evolution of Social Behavior is a 1964 scientific paper by the British evolutionary biologist W.D. Hamilton in which he lays out a kin selection. ...

The proof and discussion of its consequences however involved heavy mathematics, and was passed over by two reviewers. The third, John Maynard Smith did not completely understand it either, but recognised its significance; this passing over would later lead to friction between Hamilton and Maynard Smith, Hamilton feeling that Maynard Smith had held his work back to claim credit for the idea himself. The paper was printed in the relatively obscure Journal of Theoretical Biology, and when first published was largely ignored. The significance of it gradually increased, to the point where they are routinely cited in biology books. John Maynard Smith Professor John Maynard Smith, F.R.S. (6 January 1920 – 19 April 2004) was a British evolutionary biologist and geneticist. ... The Journal of Theoretical Biology is a scientific journal dealing with all mathematical and computational aspects of biology. ...

A large part of the discussion related to the evolution of eusociality in insects of the orders Homoptera (aphids), Isoptera (termites) and Hymenoptera (bees and wasps) based on their unusual haplodiploid sex-determination system, that meant that "super sisters" were more related to their sisters than to their own offspring and so ought to help their mother produce more sisters rather than breed themselves. However, there are later issues with this application that make it less clear-cut, such as the mothers mating with multiple males reduces relatedness between sisters. Eusociality is the phenomenon of reproductive specialisation found in some species of animal. ... Suborders Heteroptera Homoptera Hemiptera is an order of insects, comprising some 67,500 known species in two suborders, Heteroptera and Homoptera. ... Families Adelgidae Aphididae Pemphigidae Phylloxeridae and several more Aphids (superfamily Aphidoidea) are small plant-sucking insects. ... Families Mastotermitidae Kalotermitidae Termopsidae Hodotermitidae Rhinotermitidae Serritermitidae Termitidae Reference: Earthlife as of 2002-07-26 A termite (also known as a white ant) is any member of the order Isoptera, a group of social insects that eat wood and other cellulose-rich vegetable matter. ... Families Mastotermitidae Kalotermitidae Termopsidae Hodotermitidae Rhinotermitidae Serritermitidae Termitidae Reference: Earthlife as of 2002-07-26 A termite (also known as a white ant) is any member of the order Isoptera, a group of social insects that eat wood and other cellulose-rich vegetable matter. ... Suborders Apocrita Symphyta Many families, see article Hymenoptera is one of the larger orders of Insects, comprising the sawflies, wasps, bees, and ants. ... Families Andrenidae Anthophoridae Apidae Colletidae Ctenoplectridae Halictidae Heterogynaidae Megachilidae Melittidae Oxaeidae Sphecidae Stenotritidae This article is about the insect. ... Families See text. ... A haplodiploid species is one in which one of the sexes has haploid cells (cells containing one copy of each chromosome) and the other has diploid cells (cells containing two copies of each chromosome). ... A sex-determination system is a biological system that determines the development of sexual characteristics in an organism. ...

Extraordinary sex ratios

Between 1964 and 1978 Hamilton was a lecturer at Imperial College London. Whilst there he published a paper in Science on "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary" sex ratios were nearly always 1:1 (but see Edwards 1998), and likewise extraordinary sex ratios, particularly in wasps, needed explanations. Hamilton had been introduced to the idea and formulated its solution in 1960 when he had been assigned to help Fisher's pupil A.W.F. Edwards test the Fisherian sex ratio hypothesis. Hamilton combined his extensive knowledge of natural history with deep insight into the problem, opening up a whole new area of research. Imperial College London is a prestigious British academic institution focussing on science, engineering and medicine, complemented by a business school. ... Science is the journal of the American Association for the Advancement of Science (AAAS). ... Sex ratio by country for total population Sex ratio is the ratio of males to females in a population. ... Professor Anthony William Fairbank Edwards (born 1935) is a British statistician, geneticist and evolutionary biologist. ... Table of natural history, 1728 Cyclopaedia Natural history is an umbrella term for what are now usually viewed as several distinct scientific disciplines. ...

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 took his own life in poverty in 1975, and Hamilton and Maynard Smith were among the few present at the funeral. John Maynard Smith Professor John Maynard Smith, F.R.S. (6 January 1920 – 19 April 2004) was a British evolutionary biologist and geneticist. ... George R. Price (1922 - January 6, 1975) was a American population geneticist. ... In game theory, an evolutionarily stable strategy (or ESS; also evolutionary stable strategy) is a strategy which if adopted by a population cannot be invaded by any competing alternative strategy. ... Game theory is a branch of applied mathematics that studies strategic situations where players choose different actions in an attempt to maximize their returns. ... The Price equation (also known as Prices equation) is a covariance equation which is a mathematical description of evolution and natural selection. ...

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. Richard Dawkins Clinton Richard Dawkins DSc, FRS, FRSL (known as Richard Dawkins; born March 26, 1941) is an eminent British ethologist, evolutionary theorist, and popular science writer who holds the Charles Simonyi Chair in the Public Understanding of Science at Oxford University. ... The Selfish Gene is a somewhat controversial book by Richard Dawkins. ...

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. Harvard University campus (old map) Harvard University (incorporated as The President and Fellows of Harvard College) is an accredited private university in Cambridge, Massachusetts. ... The premises of the Royal Society in London (first four properties only). ... The Royal Geographical Society with the associated Institute of British Geographers is a learned society of geography and geographers. ... The University of São Paulo (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. Simultaneously, he was elected a Foreign Honorary Member of American Academy of Arts and Sciences. His arrival sparked protests and sit-ins from students who did not like his association with sociobiology. There he worked with the economist Robert Axelrod on the prisoner's dilemma. Evolutionary biology is a subfield of population biology concerned with the origin and descent of species, as well as their change over time, i. ... This article is about the University of Michigan in Ann Arbor. ... The House of the Academy, Cambridge, Massachusetts. ... A sit-in or sit-down is a form of direct action that involves one or more persons nonviolently occupying an area for protest, often political, social, or economic change. ... Sociobiology is a synthesis of scientific disciplines that attempts to explain behaviour in all species by considering the evolutionary advantages of social behaviours. ... An economist is an individual who studies, develops, and applies theories and concepts from economics, and writes about economic policy. ... This article is about a political scientist. ... Will the two prisoners cooperate to minimize total loss of liberty or will one of them, trusting the other to cooperate, betray him so as to go free? Many points in this article may be difficult to understand without a background in the elementary concepts of game theory. ...

Chasing the Red Queen

He also published the Red Queen theory of the evolution of sex. This was named for a character in Lewis Carroll's Through the Looking Glass. The Red Queens Hypothesis, Red Queen, Red Queens race or Red Queen Effect is an evolutionary hypothesis to explain the advantage of sex at the level of individuals, and the constant evolutionary arms race between competing species. ... Look up Sex in Wiktionary, the free dictionary. ... Lewis Carroll. ... Through the Looking-Glass, and What Alice Found There (1871) is a work of childrens literature by Lewis Carroll (Charles Lutwidge Dodgson), and is the sequel to Alices Adventures in Wonderland. ...

"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!" text

Hamilton hypothesised that sex had evolved because new combinations of genes could be presented to parasites - organisms with sex were able to continuously run away from their parasites. A parasite is an organism that lives in or on the living tissue of a host organism at the expense of it. ...

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 Fellowship of the Royal Society was founded in 1660. ... Sir Richard Southwood, DL, FRS, is emeritus professor of the University of Oxfords department of zoology. ... College name New College Named after Mary, mother of Jesus Established 1379 Sister College Kings College Warden Prof. ... The University of Oxford, located in the city of Oxford in England, is the oldest university in the English-speaking world. ...

From 1994 Hamilton found companionship with Maria Luisa Bozzi, an Italian science journalist and author. This article or section does not cite its references or sources. ...

On the Origin of AIDS

During the 1990s Hamilton became increasingly convinced by the controversial argument that the origin of the AIDS epidemic lay in oral polio vaccines (the OPV AIDS hypothesis) in Africa during the 1950s. Letters by Hamilton to Science were rejected by the journal, amid accusations that the medical establishment were ranging against the OPV hypothesis. AIDS (Acquired Immunodeficiency Syndrome or Acquired Immune Deficiency Syndrome, sometimes written Aids) is a human disease characterized by progressive destruction of the bodys immune system. ... Two polio vaccines are used throughout the world to combat polio. ... According to the oral polio vaccine (OPV) AIDS hypothesis, the AIDS pandemic originated from live polio vaccines prepared in chimpanzee tissue cultures (at least some of which were almost certainly contaminated with chimpanzee SIV) which were administered to up to one million Africans between 1957 and 1960. ... Science is the journal of the American Association for the Advancement of Science (AAAS). ...

To find indirect evidence of the OPV hypothesis by assessing natural levels of SIV in primates, he and two others ventured on a field trip to the war-torn Democratic Republic of the Congo, where he contracted malaria. He was rushed home and spent six weeks in hospital before dying from a cerebral haemorrhage. Malaria (Medieval Italian: mala aria — bad air) and formerly called ague or marsh fever in English, is an infectious disease which causes about 350–500 million infections in humans and approximately 1. ...

Postscript

A secular memorial service (he was an atheist) was held at the Chapel of New College, Oxford on Saturday 1st July 2000, organised by Richard Dawkins. For information about the band, see Atheist (band). ... College name New College Named after Mary, mother of Jesus Established 1379 Sister College Kings College Warden Prof. ... (Redirected from 1st July) July 1 is the 182nd day of the year (183rd in leap years) in the Gregorian Calendar, with 183 days remaining. ... This article is about the year 2000. ... Richard Dawkins Clinton Richard Dawkins DSc, FRS, FRSL (known as Richard Dawkins; born March 26, 1941) is an eminent British ethologist, evolutionary theorist, and popular science writer who holds the Charles Simonyi Chair in the Public Understanding of Science at Oxford University. ...

His body was interred in Wytham Woods, according to his request: Wytham is a small village in central Oxfordshire on the south bank of the River Thames, three miles from Oxford. ...

"- 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.

Conclusion

Hamilton is remembered as one of the greatest evolutionary theorists of the twentieth century. The other contender for the title is R.A. Fisher, and Hamilton would humbly defer to him. The concepts mentioned above are only the most important of his contributions, and many more may come to light. Sir Ronald Fisher Sir Ronald Aylmer Fisher, FRS (17 February 1890 – 29 July 1962) was a British eugenicist, evolutionary biologist, geneticist and statistician. ...

Awards

• 1978 Foreign Honorary Member of American Academy of Arts and Sciences
• 1980 Fellow of the Royal Society of London
• 1982 Newcomb Cleveland Prize of the American Association for the Advancement of Science
• 1988 Darwin Medal of the Royal Society of London
• 1989 Scientific Medal of the Linnean Society
• 1991 Frink Medal of Zoological Society of London
• 1992/3 Wander Prize of the University of Bern
• 1993 Crafoord Prize of the Royal Swedish Academy of Sciences
• 1993 Kyoto Prize of the Inamori Foundation
• 1995 Frissen Prize of the Fyssen Fondation

The House of the Academy, Cambridge, Massachusetts. ... The premises of the Royal Society in London (first four properties only). ... The American Association for the Advancement of Science (AAAS) is an organization that promotes cooperation between scientists, defends scientific freedom, encourages scientific responsibility and supports scientific education for the betterment of all humanity. ... The Darwin Medal is given by the Royal Society on even years to a biologist or a husband and wife team of biologists. ... The premises of the Royal Society in London (first four properties only). ... The Linnean Society of London is the worlds premier society for the study and dissemination about taxonomy. ... The University of Berne is a university in the Swiss capital of Berne. ... The Crafoord Prize was established in 1980 by Holger Crafoord, the inventor of the artificial kidney, and his wife Anna-Greta Crafoord. ... The Royal Swedish Academy of Sciences or , founded in 1739 by King Frederick I, is one of the Royal Academies in Sweden. ... The Kyoto Prize (京都賞) has been awarded annually since 1984 by the Inamori Foundation, founded by Kazuo Inamori (fortune from ceramics). ...

Biographies

• Alan Grafen has written a biographical memoir for the Royal Society. See http://users.ox.ac.uk/~grafen/cv/WDH_memoir.pdf
• A book is also in press: Segerstråle, U. 2005 Nature's oracle: an intellectual biography of evolutionist W. D. Hamilton. Oxford University Press.

Works

Collected Papers

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

• Hamilton, W.D. (1996) Narrow Roads of Gene Land vol. 1: Evolution of Social Behaviour Oxford University Press,Oxford. ISBN 0716745305
• Hamilton, W.D. (2002) Narrow Roads of Gene Land vol. 2: Evolution of Sex Oxford University Press,Oxford. ISBN 0198503369
• Hamilton, W.D. (in press) Narrow roads of Gene Land, vol. 3: The final years (with essays by coauthors, ed. M. Ridley). Oxford University Press, Oxford. ISBN 0198566905

Significant Papers by Hamilton

• Hamilton, W.D. (1964). The genetical evolution of social behaviour I and II. — Journal of Theoretical Biology 7: 1-16 and 17-52. pubmed I pubmed II
• Hamilton, W.D. (1967). Extraordinary sex ratios. Science 156: 477-488. pubmed JSTOR
• Hamilton, W.D. (1975). Innate social aptitudes of man: an approach from evolutionary genetics. in R. Fox (ed.), Biosocial Anthropology, Malaby Press, London, 133-53.
• Axelrod, R. and W.D. Hamilton (1981) The evolution of co-operation Science 211: 1390-6 Pubmed, JSTOR
• Hamilton, W.D. (2000) My intended burial and why, Ethology Ecology and Evolution 12 111-122 link

Science is the journal of the American Association for the Advancement of Science (AAAS). ... This article is about a political scientist. ... Science is the journal of the American Association for the Advancement of Science (AAAS). ...

Other references

• Edwards, A.W.F. (1998), Notes and Comments. Natural selection and sex ratio: Fisher's sources. American Naturalist 151: 564-569
• Fisher, R.A. (1930). The Genetical Theory of Natural Selection. Clarendon Press, Oxford.
• Ford E.B. (1945) New Naturalist 1: Butterflies. Collins: London.
• Maynard Smith, J. and G.R. Price (1973) The logic of animal conflict. Nature 146: 15—18.

Professor Anthony William Fairbank Edwards (born 1935) is a British statistician, geneticist and evolutionary biologist. ... Sir Ronald Fisher Sir Ronald Aylmer Fisher, FRS (17 February 1890 – 29 July 1962) was a British eugenicist, evolutionary biologist, geneticist and statistician. ... The Genetical Theory of Natural Selection is a book by Ronald Fisher. ... This article is about the British ecological geneticist E.B. Henry Ford. ... John Maynard Smith Professor John Maynard Smith, F.R.S. (6 January 1920 – 19 April 2004) was a British evolutionary biologist and geneticist. ... George R. Price (1922 - January 6, 1975) was a American population geneticist. ... First title page, November 4, 1869 Nature is one of the oldest and most reputable scientific journals, first published on 4 November 1869. ...

External links

• Obituaries and reminiscences
• Royal Society citation
• Truth and Science: Bill Hamiltonís 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

• W.D. Hamilton's work in game theory

AIDS

• polio vaccines and AIDS