George C. Williams (1926–2010) Incisive Thinker Who Influenced a Generation of Evolutionary Biologists

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COMMENt Obituary George C. Williams (1926–2010) Incisive thinker who influenced a generation of evolutionary biologists.

n 1978, at the age of 52, the great genetics in social behaviour, even of humans. evolutionary theorist George C. Williams And Richard Dawkins’s 1976 book The Selfish began to chronicle his own senescence, Gene popularized some of Williams’s ideas. Irecording once a year how long it took to That said, gene-level selection and inclusive run 1,700 metres round a track in Stony fitness were not universally accepted then, , Stony Brook Univ. Brook Stony , Brook, New York. Williams presented the and still meet with occasional criticism — S graph of his 12 years of slowing speed at his notably from researchers trying to explain acceptance speech for the Crafoord Prize in altruism and eusociality, for example. These

Bioscience that he shared with Ernst Mayr ideas remain, nonetheless, cornerstones of Service Media and John Maynard Smith in 1999. He later modern biological theory. published it in The Quarterly Review of Biol- ogy, with which he was involved for 32 years. competition not cooperation The plot encapsulated his lifelong fascina- Williams made further influential tion: why do we decline with age? contributions. With his 1975 book Sex and Williams died on 8 September, aged 84. Evolution, he was among the first to offer Little known to the public, this tall, reserved explanations for the puzzling prevalence of man with an Abraham Lincoln beard will sexual reproduction. He pointed out that it be remembered by evolutionary biologists is yet another example of competition, not as one of the most incisive thinkers of the cooperation, being the dominant force in twentieth century. His major contribution, evolution — with genes from each parent the theory of gene-level natural selection, battling for influence within the same left a profound and enduring stamp on fields genome. (He saw a bright future for the fields from sociobiology and evolutionary psychol- of genetic imprinting and epigenetics.) ogy to behavioural ecology. He spoke slowly Williams went even further with his reduc- and little, but when he spoke, you listened: tionist view of natural selection in Natural his words were full of insight and flashes of be expected to persist and even increase in Selection: Domains, Levels and Challenges, dry wit. abundance as long as, on balance, they boost his 1992 book about information and matter. an individual’s fitness. He also pointed out He pointed out that what is of importance in gene-level selection that selection should be weaker in older age evolution is the information that is contained After a stint in the US Army, working on a because fewer individuals are alive to be in genes, genotypes and gene pools, not the water purification plant in Italy during the subject to it — an idea for which Williams physical objects — a position reminiscent of Second World War, Williams finished his shares credit with Peter Medawar. Dawkins’s ‘meme’ concept. BA in zoology at the University of California, The dominant narrative of early 1960s Williams returned late in life to his abiding Berkeley in 1949. He got his PhD from the evolutionary biology was that natural selec- concern — ageing. University of California, Los Angeles in 1955 tion acts at the level of the group or even “He spoke slowly In 1994 he wrote the for work on the ecology of the blenny — a for ‘the good of the species’. Even death was and little, but book Why We Get type of fish. There followed a postdoc at the explained in a group-selectionist light — as when he spoke, Sick: the New Sci- University of Chicago and an assistant pro- creating space for the next generation. Wil- you listened: his ence of Darwinian fessorship at Michigan State University. In liams skewered this thinking, which he felt words were full Medicine with the 1960, Williams moved to the State University was “sloppy” and “anti-Darwinian”, in his of insight and physician Randolph of New York at Stony Brook. He later became most influential book, Adaptation and Nat- flashes of dry Nesse. Williams and one of the first professors in its newly formed ural Selection (1966). In it he proposed that wit.” Nesse proposed Department of Ecology and Evolution. There natural selection almost always acts more that disease symp- he remained until his retirement in 1990 — directly, swiftly and strongly at the level of toms should be understood, and treatment the year I arrived as assistant professor and the gene or the individual than at the level informed, by the long evolutionary history inherited his freezer of Icelandic eel samples. of the group or even species. He also railed that shaped immune responses. Their work He had spent two sabbaticals in Iceland, was against ‘pan-adaptationism’ — the idea that has spawned a new field of study, evolution- fluent in Icelandic and published on the every feature is adaptive: he showed that ary medicine. European and American species of eels and adaptations have to have fitness-enhancing It is a cruel irony that this brilliant man their potential hybrids on Iceland. effects at the level of the individual rather who first explained senescence died of In 1957, he published his seminal paper than at the level of the species. Alzheimer’s disease. ■ ‘Pleiotropy, Natural Selection, and the Adaptation and Natural Selection was Evolution of Senescence’ in the journal way ahead of its time; its impact was felt for Axel Meyer is professor of zoology and Evolution. He argued that genes that decades. Following Williams, E. O. Wilson evolutionary biology in the Department enhance fitness early in life but have det- extended gene-level and individual selec- of Biology at the University of Konstanz, rimental effects later in life — genes with tion in Sociobiology: The New Synthesis — D78457 Konstanz, Germany. ‘antagonistic pleiotropic effects’ — would his contro versial 1975 book on the role of e-mail: [email protected]