George Christopher Williams

George Christopher Williams (May 12, 1926 –

September 8, 2010) was an American evolutionary George C. Williams
biologist.[1][2][3]

Williams was a professor of biology at the State

University of New York at Stony Brook who was best
known for his vigorous critique of group selection. The
work of Williams in this area, along with W. D.
Hamilton, John Maynard Smith, Richard Dawkins, and
others led to the development of the gene-centered
view of evolution in the 1960s.

Academic work
Williams' 1957 paper Pleiotropy, Natural Selection,
and the Evolution of Senescence is one of the most
Born George Christopher Williams
influential in 20th century evolutionary biology, and
May 12, 1926
contains at least 3 foundational ideas.[4] The central
Charlotte, North Carolina, U.S.
hypothesis of antagonistic pleiotropy remains the
prevailing evolutionary explanation of senescence. In Died September 8, 2010 (aged 84)
this paper Williams was also the first to propose that Alma mater UCLA
senescence should be generally synchronized by Known for theories of natural selection
natural selection. According to this original Awards Daniel Giraud Elliot Medal (1992)
formulation
Crafoord Prize (1999)

... if the adverse genic effects appeared Scientific career

earlier in one system than any other, they Fields Biology

would be removed by selection from that Institutions Stony Brook University
system more readily than from any other.
In other words, natural selection will
always be in greatest opposition to the
decline of the most senescence-prone
system.

This important concept of synchrony of senescence was taken up a short time later by John Maynard
Smith, and the origin of the idea is often misattributed to him, including in his obituary in the journal
Nature.[5] Finally, Williams' 1957 paper was the first to outline the "grandmother hypothesis". William's
formulation stated that natural selection might select for menopause and post-reproductive life in females
(though not explicitly mentioning grandchildren or the inclusive fitness contribution of grand-parenting).
In his first book, Adaptation and Natural Selection (1966), Williams advocated a "ground rule - or
perhaps doctrine would be a better term - ... that adaptation is a special and onerous concept that should
only be used where it is really necessary",[6] and that, when it is necessary, selection among genes or
individuals would in general be the preferable explanation for it. He elaborated this view in later books
and papers, which contributed to the development of a gene-centered view of evolution. Richard Dawkins
built upon Williams' ideas around selection and genes in his book The Selfish Gene (1976).[7]

Williams was also well known for his work on the evolution of sex, and was an advocate of evolutionary
medicine.

In Sex and Evolution (1975), he attempted to explain why many species use exclusive sexual
reproduction despite its “twofold cost". He proposed several explanatory models ( “aphid-rotifer model,”
the “strawberry-coral model,” and “elm-oyster model"), though found all of them insufficient.[8] He even
considered the possibility that sex is a maladaption for some species:

When major taxonomic groups all share a certain feature, it is unlikely that the feature has
the same adaptive significance throughout the group. It may even be maladaptive for the
majority... The fact that parthenogenesis or its equivalent, if found in a vertebrate population,
has always replaced sexual reproduction entirely, is decisive evidence of the maladaptive
nature of sexuality in these organisms. (Chapter 9)

In later books, including Natural Selection: Domains, Levels and Challenges, Williams softened his
views on group selection, recognizing that clade selection, trait group selection and multilevel selection
did sometimes occur in nature, something he had earlier thought to be so unlikely it could be safely
ignored.[9][10]

Williams became convinced that the genic neo-Darwinism of his earlier years, while
essentially correct as a theory of microevolutionary change, could not account for
evolutionary phenomena over longer time scales, and was thus an "utterly inadequate
account of the evolution of the Earth's biota" (1992, p. 31). In particular, he became a staunch
advocate of clade selection – a generalisation of species selection to monophyletic clades of
any rank – which could potentially explain phenomena such as adaptive radiations, long-term
phylogenetic trends, and biases in rates of speciation/extinction. In Natural Selection (1992),
Williams argued that these phenomena cannot be explained by selectively-driven allele
substitutions within populations, the evolutionary mechanism he had originally championed
over all others. This book thus represents a substantial departure from the position of
Adaptation and Natural Selection.[11]

Academic career
Williams received a Ph.D. in biology from the University of California at Los Angeles in 1955. At Stony
Brook he taught courses in marine vertebrate zoology, and he often used ichthyological examples in his
books.
In 1992, Williams was awarded the Daniel Giraud Elliot Medal from the National Academy of
Sciences.[12] He won the Crafoord Prize for Bioscience jointly with Ernst Mayr and John Maynard Smith
in 1999. Richard Dawkins describes Williams as "one of the most respected of American evolutionary
biologists".[13]

Books
Williams, G.C. 1966. Adaptation and Natural Selection. Princeton University Press,
Princeton, N.J.
Williams, G.C., ed. 1971. Group Selection. Aldine-Atherton, Chicago.
Williams, G.C. 1975. Sex and Evolution. Princeton University Press, Princeton, N.J.
Paradis, J. and G.C. Williams. 1989. T.H. Huxley's Evolution and Ethics : with New Essays
on its Victorian and Sociobiological Context. Princeton University Press, Princeton, N.J.
Williams, G.C. 1992. Natural Selection: Domains, Levels, and Challenges. Oxford University
Press, New York.
Nesse, R.M. and G.C. Williams. 1994. Why We Get Sick : the New Science of Darwinian
Medicine. Times Books, New York.
Williams, G.C. 1996. Plan and Purpose in Nature. Weidenfeld & Nicolson, London
(published in the U.S. in 1997 as The Pony Fish's Glow : and Other Clues to Plan and
Purpose in Nature. Basic Books, New York).

Selected papers
Williams, G. C. (1957). "Pleiotropy, natural selection, and the evolution of senescence".
Evolution. 11 (4): 398–411. doi:10.2307/2406060 (https://doi.org/10.2307%2F2406060).
JSTOR 2406060 (https://www.jstor.org/stable/2406060).
Williams, G. C.; Williams, D. C. (1957). "Natural selection of individually harmful social
adaptations among sibs with special reference to social insects". Evolution. 11 (1): 32–39.
doi:10.2307/2405809 (https://doi.org/10.2307%2F2405809). JSTOR 2405809 (https://www.j
stor.org/stable/2405809).
Williams, G. C. (1966). "Natural selection, the costs of reproduction, and a refinement of
Lack's principle". The American Naturalist. 100 (916): 687–690. doi:10.1086/282461 (https://
doi.org/10.1086%2F282461). JSTOR 2459305 (https://www.jstor.org/stable/2459305).
S2CID 84993886 (https://api.semanticscholar.org/CorpusID:84993886).
Williams, G. C.; Mitton, J. B. (1973). "Why reproduce sexually?". Journal of Theoretical
Biology. 39 (3): 545–554. doi:10.1016/0022-5193(73)90067-2 (https://doi.org/10.1016%2F0
022-5193%2873%2990067-2). PMID 4730017 (https://pubmed.ncbi.nlm.nih.gov/4730017).
Williams, G. C. (1979). "The question of adaptive sex ratio in outcrossed vertebrates".
Proceedings of the Royal Society B: Biological Sciences. 205 (1161): 567–580.
Bibcode:1979RSPSB.205..567W (https://ui.adsabs.harvard.edu/abs/1979RSPSB.205..567
W). doi:10.1098/rspb.1979.0085 (https://doi.org/10.1098%2Frspb.1979.0085).
JSTOR 77446 (https://www.jstor.org/stable/77446). PMID 42061 (https://pubmed.ncbi.nlm.ni
h.gov/42061). S2CID 35775599 (https://api.semanticscholar.org/CorpusID:35775599).
Hrdy, S. B.; Williams, G. C. (1983). "Behavioral biology and double standard". In Wasser, S.
K. (ed.). Social Behavior of Female Vertebrates. New York, United States: Academic Press.
pp. 3–17. ISBN 9780124313415.
Taylor, P. O.; Williams, G. C. (1984). "Demographic parameters at evolutionary equilibrium".
Canadian Journal of Zoology. 62 (11): 2264–2271. doi:10.1139/z84-329 (https://doi.org/10.1
139%2Fz84-329).
 Williams, G. C. (1985). "A defense of reductionism in evolutionary biology". In Dawkins, R.;
Ridley, M. (eds.). Oxford Surveys in Evolutionary Biology: Volume 2. Oxford, United
Kingdom: Oxford University Press. pp. 1–27. ISBN 9780198541745.
Williams, G. C. (1988). "Huxley's Evolution and Ethics in Sociobiological Perspective".
Zygon. 23 (4): 383–438. doi:10.1111/j.1467-9744.1988.tb00852.x (https://doi.org/10.1111%
2Fj.1467-9744.1988.tb00852.x).
Williams, G. C.; Nesse, R. M. (1991). "The dawn of Darwinian medicine". The Quarterly
Review of Biology. 66 (1): 1–22. doi:10.1086/417048 (https://doi.org/10.1086%2F417048).
JSTOR 2830330 (https://www.jstor.org/stable/2830330). PMID 2052670 (https://pubmed.ncb
i.nlm.nih.gov/2052670). S2CID 40357032 (https://api.semanticscholar.org/CorpusID:403570
32).
Williams, G. C. (1992). "Gaia, nature worship and biocentric fallacies". The Quarterly
Review of Biology. 67 (4): 479–486. doi:10.1086/417796 (https://doi.org/10.1086%2F41779
6). JSTOR 2832018 (https://www.jstor.org/stable/2832018). S2CID 84360055 (https://api.se
manticscholar.org/CorpusID:84360055).
Williams, G. C. (1993). "Mother nature is a wicked old witch" (http://www.sunypress.edu/p-1
631-evolutionary-ethics.aspx). In Nitecki, M. H.; Nitecki, D. V. (eds.). Evolutionary Ethics.
Albany, United States: SUNY Press. pp. 217–232. ISBN 9780791414996.
Williams, G. C. (1999). "The Tithonus error in modern gerontology". The Quarterly Review of
Biology. 74 (4): 405–415. doi:10.1086/394111 (https://doi.org/10.1086%2F394111).
JSTOR 2664720 (https://www.jstor.org/stable/2664720). PMID 10672642 (https://pubmed.nc
bi.nlm.nih.gov/10672642). S2CID 83654243 (https://api.semanticscholar.org/CorpusID:8365
4243).
Williams, G. C. (2008). "A package of information" (http://www.edge.org/documents/ThirdCul
ture/h-Ch.1.html). In Brockman, J. (ed.). The Third Culture: Beyond The Scientific
Revolution. New York, United States: Touchstone. pp. 38–50. ISBN 9780684823447.
Williams, G. C. (2008). "Darwinian medicine". Encyclopedia of Life Sciences (ELS).
Chichester, United Kingdom: John Wiley & Sons.
doi:10.1002/9780470015902.a0001726.pub2 (https://doi.org/10.1002%2F9780470015902.a
0001726.pub2). ISBN 978-0470016176.

Controversy
Williams supervised an undergraduate project in 1985 which consisted of a student, Mitchell Behm,
tossing live animals into tubs with domesticated ferrets, which Behm subsequently admitted he partly did
"for his own amusement." Dr. Charles Middleton, Director of the Division of Laboratory Animal
Resources stated, "If animals are just going to tear each other up, the experiment would not have been
approved." Ferrets were illegal in New York at the time, without a license, which neither individual had.

After police investigation, Williams received a formal reprimand from SUNY Stony Brook for never
receiving approval because of not detailing the pain of the animals involved, and for allowing non-
campus animals to participate. Because the statute of limitations had expired, Williams narrowly escaped
strict disciplinary action, in addition to criminal prosecution. Dr. Mark Lerman, Medical Director of
Lifeline for Wildlife said there was no justification and that the experiment was completely useless.[14]
The experiments were in direct violation of the Animal Welfare Act of 1966[15] and its PHS policy
amendment introducing the Institutional Animal Care and Use Committee model,[16] to which SUNY and
all of its researchers were subject in 1988.

References
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1181013/http://old.richarddawkins.net/articles/527711-george-c-williams-1926-2010).
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926-2010) on January 21, 2014. Retrieved September 10, 2010.
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010/26821/). September 9, 2010.
3. Meyer, A. (2010). "George C. Williams (1926–2010)" (https://doi.org/10.1038%2F467790a).
Nature. 467 (7317): 790. Bibcode:2010Natur.467..790M (https://ui.adsabs.harvard.edu/abs/
2010Natur.467..790M). doi:10.1038/467790a (https://doi.org/10.1038%2F467790a).
PMID 20944730 (https://pubmed.ncbi.nlm.nih.gov/20944730).
4. Williams, George C. (1957). "Pleiotropy, Natural Selection, and the Evolution of
Senescence". Evolution. 11 (4): 398–411. doi:10.2307/2406060 (https://doi.org/10.2307%2F
2406060). JSTOR 2406060 (https://www.jstor.org/stable/2406060).
5. Szathmáry, E. R.; Hammerstein, P. (2004). "Obituary: John Maynard Smith (1920–2004)".
Nature. 429 (6989): 258–259. Bibcode:2004Natur.429..258S (https://ui.adsabs.harvard.edu/
abs/2004Natur.429..258S). doi:10.1038/429258a (https://doi.org/10.1038%2F429258a).
PMID 15152239 (https://pubmed.ncbi.nlm.nih.gov/15152239).
6. Adaptation and Natural Selection (https://www.amazon.com/dp/0691026157) p4
7. Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way We
Think (https://archive.org/details/richarddawkinsho00alan/page/67). New York, New York:
Oxford University Press. p. 67 (https://archive.org/details/richarddawkinsho00alan/page/67).
ISBN 978-0-19-929116-8.
8. Dawkins, Richard (October 2010). "George C. Williams (1926–2010)" (https://www.science.o
rg/doi/10.1126/science.1197701). Science. 330 (6000): 49–49.
doi:10.1126/science.1197701 (https://doi.org/10.1126%2Fscience.1197701). ISSN 0036-
8075 (https://search.worldcat.org/issn/0036-8075).
9. Brockman, John. "Part One: The Evolutionary Idea" (http://edge.org/conversation/part-oneth
e-evolutionary-idea). Retrieved September 10, 2013.
10. George C. Williams, Natural Selection: Domains, Levels and Challenges, (Oxford University
Press, 1992), 23-55
11. Okasha, Samir. "Maynard Smith on the levels of selection question" (https://web.archive.org/
web/20100605080942/http://www.asu.edu/clas/csdc/events/pdf/Okasha%20Levels%20of%2
0Selection.pdf) (PDF). Archived from the original (http://www.asu.edu/clas/csdc/events/pdf/
Okasha%20Levels%20of%20Selection.pdf) (PDF) on June 5, 2010. Retrieved
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ine.org/site/PageServer?pagename=AWARDS_elliot). National Academy of Sciences.
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14. "Investigation Continues on Animal Abuse" (https://dspace.sunyconnect.suny.edu/bitstream/
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15. "USDA Animal Care: Animal Welfare Act and Animal Welfare Regulations" (https://www.aphi
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laws/phs-policy.htm). National Institutes of Health: Office of Laboratory Animal Welfare.
Retrieved December 22, 2021.

External links
Official website (https://web.archive.org/web/20090314001933/http://life.bio.sunysb.edu/ee/
people/williamsindex.html)
Article from Science (https://www.science.org/doi/full/10.1126/science.304.5675.1235) by
Carl Zimmer
A Conversation With George C. Williams by Frans Roes (http://www.stephenjaygould.org/lib
rary/williams_interview.html) Archived (https://web.archive.org/web/20190726054911/http://
www.stephenjaygould.org/library/williams_interview.html) July 26, 2019, at the Wayback
Machine
Obituary by Richard Dawkins, October 1, 2010 (http://richarddawkins.net/articles/527711-ge
orge-c-williams-1926%E2%80%932010) Archived (https://web.archive.org/web/2010100417
2157/http://richarddawkins.net/articles/527711-george-c-williams-1926%E2%80%932010)
October 4, 2010, at the Wayback Machine
Stephen C. Stearns, "George Christopher Williams", Biographical Memoirs of the National
Academy of Sciences (2011) (http://www.nasonline.org/publications/biographical-memoirs/m
emoir-pdfs/williams-george.pdf)

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