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Richard Lewontin (1929–2021)

Lewontin was an evolutionary biologist, population geneticist, and public intellectual whose work moved between mathematical genetics, the critique of adaptationism, and the politics of science. His formalisation of the conditions for natural selection (1970) gave evolutionary biology one of its clearest theoretical statements. His critique of the adaptationist programme — with Gould, in the “Spandrels” paper (1979) — challenged the gene-centric mainstream and shaped a generation of argument about what natural selection can and cannot explain. His broader programme, developed with Richard Levins, insisted that organisms and environments are mutually constitutive, not independent — a position that anticipated niche construction theory by nearly two decades.

Life

Born 29 March 1929 in New York City. BA from Harvard (1951). MSc in mathematical statistics from Columbia (1952). PhD in zoology from Columbia (1954) under Theodosius Dobzhansky, working on Drosophila population genetics. Faculty at North Carolina State University, then the University of Rochester, then the University of Chicago (1965–73), then Harvard (Alexander Agassiz Professor of Zoology, 1973–98; emeritus thereafter). Also held an appointment at the Museum of Comparative Zoology.

Politically active throughout his career — a Marxist who understood the politics of science as inseparable from its content. Member of Science for the People. Co-author with Levins of The Dialectical Biologist (1985), which made the political and philosophical commitments explicit.

Died 4 July 2021, aged ninety-two.

The three-conditions formalisation

Lewontin’s 1970 paper “The units of selection” gave natural selection its most compact formal statement: evolution by natural selection occurs in any population where three conditions hold simultaneously — phenotypic variation among individuals, differential fitness correlated with that variation, and heritability of fitness. The formalisation makes clear that natural selection is not a force acting on organisms but a statistical consequence of the three conditions holding together.

The formulation is substrate-neutral in principle — it says nothing about genes, DNA, or biology specifically. Whether this generality is a strength (making Darwinian logic applicable beyond biology) or a weakness (abstracting away the biology that makes evolution what it is) became a contested question that Lewontin himself came down firmly on the restrictive side of: extending Darwinian logic beyond biology, he argued, conflates the mechanism with a metaphor.

The Spandrels critique

Gould and Lewontin, “The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptationist programme” (Proceedings of the Royal Society of London B, 1979). The most influential critique of adaptationism in twentieth-century evolutionary biology.

The argument: evolutionary biologists systematically overestimate the power of natural selection by assuming that every trait is an adaptation — shaped by selection for its current function. The adaptationist programme treats the organism as a collection of independently optimised traits, each with its own selective story, and neglects structural, developmental, and historical constraints on form. Many features of organisms are not adaptations at all but by-products of other features (spandrels), consequences of developmental architecture, or products of historical contingency.

The paper’s architectural metaphor — the spandrels of San Marco are structural by-products of the decision to mount a dome on arches, not independently designed features — became one of biology’s most cited images. The critique was broader than gene-centrism but became entangled with the gene-centric debate: Dawkins’ programme was one of its primary targets, though Gould and Lewontin aimed at the adaptationist stance across all its forms.

The Spandrels paper remains contested. Defenders of the adaptationist programme argue that Gould and Lewontin caricatured their opponents, that adaptationist reasoning is a productive heuristic even when it sometimes fails, and that the alternative — invoking constraint without testing for it — is less productive than the programme it criticises.

The organism-environment dialectic

Lewontin’s broader theoretical programme, developed with Richard Levins across several decades and articulated most fully in The Dialectical Biologist (1985) and Biology Under the Influence (2007).

The central claim: organisms do not passively adapt to pre-existing environments. Organisms construct their environments — they select habitats, modify physical conditions, transform the chemical environment, and alter the selective pressures that act on themselves and on other species. The organism and its environment are mutually constitutive; neither exists independently of the other.

This is a direct challenge to the standard picture in which the environment is given and the organism adapts to it. Lewontin argued that the standard picture smuggles in a pre-Darwinian metaphysics — the environment as a fixed stage on which organisms perform — that Darwin’s own thinking should have made untenable.

The position anticipated niche construction theory (Odling-Smee, Laland, Feldman, 2003) by nearly two decades. Lewontin’s framing was philosophical and dialectical; niche construction theory gave it a formal and empirical programme.

Molecular population genetics

Lewontin’s earlier work, before the critique became his public identity, was foundational in molecular population genetics. With Jack Hubby, he published the first gel electrophoresis survey of protein variation in natural populations of Drosophila (1966) — revealing far more genetic variation than the prevailing theory predicted. The finding that natural populations carry extensive hidden genetic variation was a major empirical discovery and contributed to the neutralist-selectionist debate: Motoo Kimura’s neutral theory (1968) proposed that most of this variation is selectively neutral, maintained by mutation-drift balance rather than by selection. Lewontin’s data fuelled the debate without settling it.

The politics of biology

Lewontin was among the most prominent scientist-critics of biological determinism. With Gould, Leon Kamin, and others in the Sociobiology Study Group, he publicly challenged E. O. Wilson’s Sociobiology (1975) on the grounds that its extension to human behaviour amounted to genetic determinism with political consequences. Not in Our Genes (1984, with Steven Rose and Leon Kamin) developed the critique at book length.

Wilson and other sociobiologists rejected the determinism characterisation, framing the programme as a research project on the evolutionary origins of behaviour rather than a political claim about its inevitability. Whether the critique’s framing was fair has been debated since.

Lewontin’s position was not that biology is irrelevant to human behaviour but that the reductionist move — from genes to traits to social arrangements — systematically obscures the role of environment, development, and social organisation. His Marxism was explicit: he understood science as situated within social relations, and the claim that human behaviour is genetically determined as serving particular political interests.

Where Lewontin stops

Lewontin’s programme is critical and dialectical. It identifies what reductionist accounts leave out — what adaptationism overlooks, what gene-centrism abstracts away, what the standard environment-as-given picture conceals — and names the alternative structure: the organism-environment dialectic, the mutual constitution of organism and world. The formal and empirical apparatus for that alternative sits outside Lewontin’s own work. Niche construction theory later supplied some of it; developmental systems theory supplied more. Lewontin’s territory is the diagnostic move and the philosophical framing, not the constructive programme that follows.

Key works

See also: Darwinism · Darwin · Dawkins · Mayr · Fisher