Wikipedia

W. D. Hamilton

(Redirected fromW.D. Hamilton)

William Donald HamiltonFRS(1 August 1936 – 7 March 2000) was a Britishevolutionary biologist,recognised as one of the most significantevolutionarytheorists of the 20th century.[1][2]Hamilton became known for histheoreticalwork expounding a rigorousgeneticbasis for the existence ofaltruism,an insight that was a key part of the development of thegene-centered view of evolution.He is considered one of the forerunners ofsociobiology.Hamilton published important work onsex ratiosand theevolution of sex.From 1984 to his death in 2000, he was aRoyal SocietyResearch Professor atOxford University.

W. D. Hamilton
Born
William Donald Hamilton

(1936-08-01)1 August 1936
Cairo,Egypt
Died7 March 2000(2000-03-07)(aged 63)
Fitzrovia,London,United Kingdom
NationalityBritish
EducationSt. John's College, Cambridge
London School of Economics
University College London
Known forKin selection,Hamilton's rule
SpouseChristine Friess
ChildrenThree daughters
Parent(s)Archibald Milne Hamiltonand Bettina Matraves Hamilton née Collier
AwardsNewcomb Cleveland Prize(1981)
Linnean Medal(1989)
Kyoto Prize(1993)
Crafoord Prize(1993)
Sewall Wright Award(1998)
Scientific career
FieldsEvolutionary biology
InstitutionsImperial College London,University of Michigan,University of Oxford
Academic advisorsJohn Hajnal
Cedric Smith
Doctoral studentsLaurence Hurst
Olivia Judson

Richard Dawkinshas written that Hamilton was "the greatest Darwinian of my lifetime".[3]

Early life

edit

Hamilton was born in 1936 inCairo,Egypt,the second of seven children. His parents were fromNew Zealand;his fatherA.M. Hamiltonwas an engineer, and his mother B.M. Hamilton was aphysician.The Hamilton family settled inKent.During theSecond World War,Hamilton was evacuated toEdinburgh.He became interested in natural history at an early age and spent his spare time collectingbutterfliesand other insects. In 1946, he discoveredE.B. Ford'sNew NaturalistbookButterflies,which introduced him to the principles of evolution bynatural selection,genetics,andpopulation genetics.

He was educated atTonbridge School,where he was in Smythe House. As a 12-year-old, he was seriously injured while playing withexplosiveshis father had that were left over from makinghand grenadesfor theHome GuardduringWorld War II.Hamilton had to have athoracotomyand parts of fingers on his right hand had to be amputated inKing's College Hospitalto save his life. He was left with scarring and needed six months to recover.

Before going up to theUniversity of Cambridge,he travelled in France and completed two years ofnational service.As anundergraduateatSt. John's Collegein Biology, he was uninspired by the "many biologists [who] hardly seemed to believe in evolution".

Hamilton's rule

edit
Main articles:Kin selectionandSocial Bonding and Nurture Kinship § Evolutionary biology theory of social behavior

Hamilton enrolled in anMSccourse in demography at theLondon School of Economics(LSE), underNorman Carrier,who helped secure grants for his studies. Later, when his work became more mathematical and genetical, he had his supervision transferred toJohn Hajnalof the LSE andCedric SmithofUniversity College London(UCL).

Both Fisher andJ. B. S. Haldanehad 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 wasSewall Wright'scoefficient 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:

whereCis the cost in fitness to the actor,rthe genetic relatedness between the actor and the recipient, andBis the fitness benefit to the recipient. Fitness costs and benefits are measured infecundity.ris a number between 0 and 1. His two 1964 papers entitledThe Genetical Evolution of Social Behaviourare now widely referenced.[4]

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 Maynard 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 theJournal of Theoretical Biologyand, when first published, was largely ignored. Recognition of its significance gradually increased to the point that it is now routinely cited in biology books.

Much of the discussion relates to the evolution ofeusocialityin insects of the orderHymenoptera(ants,beesandwasps) based on their unusualhaplodiploid 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.

The supergenes notion (sometimes called theGreen-beard effect) - that organisms may evolve genes that are able to identify identical copies in others and preferentially direct social behaviours towards them - was theoretically clarified and withdrawn by Hamilton in 1987.[5]

Spiteful behaviour

edit
Main article:Hamiltonian spite

In his 1970 paperSelfish and Spiteful Behaviour in an Evolutionary ModelHamilton 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

edit

Between 1964 and 1977, Hamilton was a lecturer atImperial College London(includingSilwood Park,where a building is named in his honour).[6]Whilst there he published a paper inScienceon "extraordinary sex ratios". Fisher (1930) had proposed a model as to why "ordinary"sex ratioswere 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 pupilA.W.F. Edwardstest the Fisherian sex ratio hypothesis. Hamilton combined his extensive knowledge ofnatural historywith deep insight into the problem, opening up a whole new area of research.

The paper introduced the concept of the "unbeatable strategy", whichJohn Maynard SmithandGeorge R. Pricewere to develop into theevolutionarily stable strategy(ESS), a concept ingame theorynot limited to evolutionary biology. Price had originally come to Hamilton after deriving thePrice 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.[7]

Hamilton was a visiting professor atHarvard Universityand later spent nine months with theRoyal Society's and theRoyal Geographical Society'sXavantina-Cachimbo Expeditionas a visiting professor at theUniversity of São Paulo.From 1978 Hamilton was Professor ofEvolutionary Biologyat theUniversity of Michigan.Simultaneously, he was elected a Foreign Honorary Member ofAmerican Academy of Arts and Sciences.[8]His arrival sparked protests andsit-insfrom students who did not like his association withsociobiology.There he worked with thepolitical scientistRobert Axelrodon theprisoner's dilemma,and was a member of the BACH group with original membersArthur Burks,Robert Axelrod,Michael Cohen,andJohn Holland.[9]

Hamilton was regarded as a poor lecturer. This shortcoming would not affect the recognition of his work, however, as it was popularised byRichard Dawkinsin the bookThe Selfish Genepublished in 1976.

Chasing the Red Queen

edit

Hamilton was an early proponent of theRed Queentheory of theevolution of sex[10](separate from the other theory of the same name previously proposed byLeigh Van Valen). This was named for a character inLewis Carroll'sThrough the Looking-Glass,who is continuously running but never actually travels any distance:

"Well, in our country," said Alice, still panting a little, "you'd generally get to somewhere else—if you ran very fast for a long time, as we've been doing."
"A slow sort of country!" said the Queen. "Now, here, you see, it takes all the running you can do, to keep in the same place. If you want to get somewhere else, you must run at least twice as fast as that!"(Carroll, pp. 46)

This theory hypothesizes that sex evolved because new and unfamiliar combinations of genes could be presented toparasites,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

edit

In 1980, he was elected aFellow of the Royal Society,and in 1984, he was invited byRichard Southwoodto be the Royal Society Research Professor in the Department of Zoology atOxford,and a fellow ofNew College,where he remained until his death.

His collected papers, entitledNarrow Roads of Gene Land,began to be published in 1996. The first volume was entitledEvolution of Social Behaviour.

Social evolution

edit

The field ofsocial evolution,of whichHamilton's Rulehas central importance, is broadly defined as being the study of the evolution of social behaviours, i.e. those that impact on the fitness of individuals other than the actor. Social behaviours can be categorized according to the fitness consequences they entail for the actor and recipient. A behaviour that increases the direct fitness of the actor is mutually beneficial if the recipient also benefits, and selfish if the recipient suffers a loss. A behaviour that reduces the fitness of the actor is altruistic if the recipient benefits, and spiteful if the recipient suffers a loss. This classification was first proposed by Hamilton in 1964.[citation needed]

Hamilton also proposed the coevolution theory ofautumn leaf coloras an example of evolutionarysignalling theory.[11]

Origin of HIV

edit

During the 1990s, Hamilton became interested in thenow-discredited hypothesisthat the origin ofHIVlay inHilary Koprowski's oralpolio vaccinetrials in Africa during the 1950s. Hamilton's letter on the topic toSciencejournal was rejected in 1996. Despite this, he spoke to the BBC supporting the hypothesis,[12]and wrote the foreword of Edward Hooper's 1999 bookThe River.To look for evidence of the hypothesis, Hamilton went on a 2000 field trip to theDemocratic Republic of the Congoto assess natural levels ofsimian immunodeficiency virusin primates.[13][14]None of the over 60 urine and faecal samples contained detectableSIV virus.[15]

Death

edit

Hamilton returned to London from Africa on 29 January 2000. He was admitted toUniversity College Hospital,London, on 30 January 2000. He was transferred toMiddlesex Hospitalon 5 February 2000 and died there on 7 March 2000. An inquest was held on 10 May 2000 at WestminsterCoroner's Courtto 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 aduodenaldiverticulumandarterialbleed through amucosal ulcer".Following reports attributing his death to complications arising frommalaria,theBBCEditorial Complaints Unit's investigation established that he had contracted malaria during his final African expedition. However, the pathologist had suggested the possibility that the ulceration and consequent haemorrhage had resulted from a pill (which might have been taken because of malarial symptoms) lodging in the diverticulum; but, even if this suggestion were correct, the link between malaria and the observed causes of death would be entirely indirect.[16]

A secular memorial service (he was anagnostic[17]) was held at the chapel ofNew College, Oxfordon 1 July 2000, organised by Richard Dawkins. He was buried nearWytham Woods.He, however, had written an essay onMy intended burial and whyin which he wrote:[18]

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 greatCoprophanaeusbeetle 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-fusedelytrawhich we will all hold over our backs. So finally I too will shine like a violet ground beetle under a stone.

The second volume of his collected papers,Evolution of Sex,was published in 2002, and the third and final volume,Last Words,in 2005.

In 1966, he married Christine Friess; the couple had three daughters, Helen, Ruth, and Rowena.[19]They amicably separated 26 years later.[citation needed]From 1994, Hamilton found companionship with Maria Luisa Bozzi, an Italian science journalist and author.[19]

Awards

edit

Biographies

edit
  • Alan Grafenhas written a biographical memoir for the Royal Society.[19]
  • A biographical book has also been published byUllica Segerstråle:Segerstråle, U. 2013.Nature's oracle: the life and work of W. D. Hamilton.Oxford University Press.ISBN978-0-19-860728-1

Works

edit

Collected papers

edit

Hamilton started to publish his collected papers 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

edit

Notes

edit
  1. ^"Obituary by Richard DawkinsArchived4 March 2016 at theWayback Machine",The Independent,10 March 2000. See also his eulogy byRichard Dawkinsreprinted in his bookA Devil's Chaplain(2003).
  2. ^BBC Radio 4 –Great Lives– 2 Feb 2010
  3. ^Dawkins, Richard (2022) [2021].Flights of Fancy: Defying Gravity by Design and Evolution.London:Head of Zeus.p. 210.ISBN978-1838937850.
  4. ^Aaen-Stockdale, C. (2017), "Selfish Memes: An Update of Richard Dawkins' Bibliometric Analysis of Key Papers in Sociobiology",Publications,5(2): 12,doi:10.3390/publications5020012,hdl:11250/2443958
  5. ^Hamilton, William D. 1987. Discriminating nepotism: expectable, common and overlooked. In Kin recognition in animals, edited by D. J. C. Fletcher and C. D. Michener. New York: Wiley.
  6. ^"WD Hamilton".TheGuardian.com.9 March 2000.
  7. ^Brown, Andrew (2000).The Darwin Wars: The Scientific Battle for the Soul of Man.London: Touchstone.ISBN978-0-684-85145-7.
  8. ^ab"William Donald Hamilton".American Academy of Arts & Sciences.Retrieved2 December2021.
  9. ^"History".
  10. ^The Red Queen Hypothesisat Indiana University. Quote: "W. D. Hamilton and John Jaenike were among the earliest pioneers of the idea."
  11. ^Hamilton, WD; Brown, SP (July 2001)."Autumn tree colours as a handicap signal".Proc. R. Soc. B.268(1475): 1489–1493.doi:10.1098/rspb.2001.1672.ISSN0962-8452.PMC1088768.PMID11454293.
  12. ^"'Scientists started Aids epidemic'".BBC News.1 September 1999.Retrieved1 September2020.
  13. ^"The Politics of a Scientific Meeting: the Origin-of-AIDS Debate at the Royal Society".Politics and the Life Sciences.20(20). September 2001.Retrieved1 September2020.
  14. ^Allen, Kate (29 April 2023)."He nearly died pursuing HIV's origins. Then this Canadian scientist set his sights on the COVID lab leak theory. Here's what he found".Toronto Star.Retrieved15 September2023.
  15. ^Horton, Richard (2000). "New data challenge OPV theory of AIDS origin".The Lancet.356(9234): 1005.doi:10.1016/S0140-6736(00)02698-2.
  16. ^"ECU Ruling: Great Lives, BBC Radio 4, 2 February 2010".BBC.Retrieved24 June2011.
  17. ^Ullica Segerstrale (28 February 2013).Nature's Oracle: The Life and Work of W.D.Hamilton.OUP Oxford. pp. 383–.ISBN978-0-19-164277-7.
  18. ^Hamilton, W. D. (2000)."My intended burial and why".Ethology Ecology and Evolution.12(2): 111–122.doi:10.1080/08927014.2000.9522807.S2CID84908650.[permanent dead link‍]
  19. ^abcdGrafen, A.(2004)."William Donald Hamilton. 1 August 1936 -- 7 March 2000"(PDF).Biographical Memoirs of Fellows of the Royal Society.50:109–132.doi:10.1098/rsbm.2004.0009.S2CID56905497.
  20. ^"William D Hamilton".Crafoord Prize.22 August 2022.Retrieved24 February2024.
  21. ^"APS Member History".search.amphilsoc.org.Retrieved2 December2021.

References

edit
edit
Wikiquote has quotations related toW. D. Hamilton.
Retrieved from "https://en.wikipedia.org/w/index.php?title=W._D._Hamilton&oldid=1259651791"