|Born||Richard Charles Lewontin
29 March 1929
New York City, New York, U.S.
North Carolina State University
University of Rochester
University of Chicago
|Alma mater||Columbia University|
|Thesis||The Effects of Population Density and Composition on Viability in Drosophila melanogaster (1955)|
|Academic advisors||Theodosius Dobzhansky|
|Doctoral students||Joseph Felsenstein
|Known for||Evolutionary biology
|Notable awards||Sewall Wright Award (1994), Crafoord Prize (2015)|
Richard Charles "Dick" Lewontin (born March 29, 1929) is an American evolutionary biologist, geneticist, academic and social commentator. A leader in developing the mathematical basis of population genetics and evolutionary theory, he pioneered the application of techniques from molecular biology, such as gel electrophoresis, to questions of genetic variation and evolution.
In a pair of seminal 1966 papers co-authored with J.L. Hubby in the journal Genetics, Lewontin helped set the stage for the modern field of molecular evolution. In 1979 he and Stephen Jay Gould introduced the term "spandrel" into evolutionary theory. From 1973 to 1998, he held an endowed chair in zoology and biology at Harvard University, and since 2003 has been a research professor there.
- 1 Early life and education
- 2 Career
- 3 Personal life
- 4 Recognition
- 5 Bibliography
- 6 References
- 7 See also
- 8 Further reading
- 9 External links
Early life and education
Lewontin was born in New York City to parents descended from late 19th-century Eastern European Jewish immigrants. He attended Forest Hills High School and the École Libre des Hautes Études in New York. In 1951 he graduated from Harvard College (BS, biology). In 1952, Lewontin received a master's degree in mathematical statistics, followed by a doctorate in zoology in 1954, both from Columbia University, where he was a student of Theodosius Dobzhansky.
He held faculty positions at North Carolina State University, the University of Rochester, and the University of Chicago. In 1973 Lewontin was appointed as Alexander Agassiz Professor of Zoology and Professor of Biology at Harvard University, holding the position until 1998.
Work in population genetics
Lewontin has worked in both theoretical and experimental population genetics. A hallmark of his work has been an interest in new technology. He was the first person to do a computer simulation of the behavior of a single locus (previous simulation work having been of models with multiple loci). In 1960 he and Ken-Ichi Kojima were the first population geneticists to give the equations for change of haplotype frequencies with interacting natural selection at two loci. This set off a wave of theoretical work on two-locus selection in the 1960s and 1970s. Their paper gave a theoretical derivation of the equilibria expected, and also investigated the dynamics of the model by computer iteration. Lewontin later introduced the D' measure of linkage disequilibrium. (An achievement that he would be less happy to claim is that he introduced the name "linkage disequilibrium" itself, one about which many population geneticists have been unenthusiastic).
In 1966, he and Jack Hubby published a paper that revolutionized population genetics. They used protein gel electrophoresis to survey dozens of loci in Drosophila pseudoobscura, and reported that a large fraction of the loci were polymorphic, and that at the average locus there was about a 15% chance that the individual was heterozygous. (Harry Harris reported similar results for humans at about the same time.) Previous work with gel electrophoresis had been reports of variation in single loci and did not give any sense of how common variation was.
Lewontin and Hubby's paper also discussed the possible explanation of the high levels of variability by either balancing selection or neutral mutation. Although they did not commit themselves to advocating neutrality, this was the first clear statement of the neutral theory for levels of variability within species. Lewontin and Hubby's paper had great impact—the discovery of high levels of molecular variability gave population geneticists ample material to work on, and gave them access to variation at single loci. The possible theoretical explanations of this rampant polymorphism became the focus of most population genetics work thereafter. Martin Kreitman was later to do a pioneering survey of population-level variability in DNA sequences while a Ph.D. student in Lewontin's lab.
Work on human genetic diversity
In a landmark paper, in 1972 Lewontin identified that most of the variation (80–85%) within human populations is found within local geographic groups and differences attributable to traditional "race" groups are a minor part of human genetic variability (1–15%). In a 2003 paper, A.W.F. Edwards criticized Lewontin's conclusion that race is an invalid taxonomic construct, terming it Lewontin's fallacy. He argued that the probability of racial misclassification of an individual based on variation in a single genetic locus is approximately 30% and the misclassification probability becomes close to zero if enough loci are studied. 
Critique of mainstream evolutionary biology
In 1975, when E. O. Wilson's book Sociobiology proposed evolutionary explanations for human social behaviors, a number of biologists responded negatively, including Lewontin, Stephen Jay Gould, Ruth Hubbard, and others.
Lewontin and his Harvard colleague Stephen Jay Gould introduced the term spandrel to evolutionary biology, inspired by the architectural term "spandrel", in an influential 1979 paper, "The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme." "Spandrels" were described as features of an organism that exist as a necessary consequence of other (perhaps adaptive) features, but do not directly improve fitness (and thus are not necessarily adaptive). The relative frequency of spandrels versus adaptations continues to stir controversy in evolutionary biology.
Lewontin was an early proponent of a hierarchy of levels of selection in his article, "The Units of Selection". He has been a major influence on philosophers of biology, notably William C. Wimsatt (who taught with Lewontin and Richard Levins at the University of Chicago), Robert Brandon and Elisabeth Lloyd (who studied with Lewontin as graduate students), Philip Kitcher, Elliott Sober, and Sahotra Sarkar. Lewontin briefly argued for the historical nature of biological causality in "Is Nature Probable or Capricious?".
In "Organism and Environment" in Scientia, and in more popular form in the last chapter of Biology as Ideology, Lewontin argued that while traditional Darwinism has portrayed the organism as a passive recipient of environmental influences, a correct understanding should emphasize the organism as an active constructor of its own environment. Niches are not pre-formed, empty receptacles into which organisms are inserted, but are defined and created by organisms. The organism-environment relationship is reciprocal and dialectical. M.W. Feldman, K.N. Laland, and F.J. Odling-Smee,  among others, have developed Lewontin's conception in more detailed models.
In the adaptationist view of evolution, the organism is a function of both the organism and environment, while the environment is only a function of itself. The environment is seen as autonomous and unshaped by the organism. Lewontin instead believed in a constructivist view, in which the organism is a function of the organism and environment, with the environment being a function of the organism and environment as well. This means that the organism shapes the environment as the environment shapes the organism. The organism shapes the environment for future generations.
Lewontin has long been a critic of traditional neo-Darwinian approaches to adaptation. In his article "Adaptation" in the Italian Enciclopedia Einaudi, and in a toned-down version in Scientific American, he emphasized the need to give an engineering characterization of adaptation separate from measurement of number of offspring, rather than simply assuming organs or organisms are at adaptive optima. Lewontin has said that his more general, technical criticism of adaptationism grew out of his recognition that the fallacies of sociobiology reflect fundamentally flawed assumptions of adaptiveness of all traits in much of the modern evolutionary synthesis.
Sociobiology and evolutionary psychology
Along with others, such as Gould, Lewontin has been a persistent critic of some themes in neo-Darwinism. Specifically, he has criticised proponents of sociobiology and evolutionary psychology such as Edward O. Wilson and Richard Dawkins, who attempt to explain animal behaviour and social structures in terms of evolutionary advantage or strategy. He and others criticize this approach when applied to humans, as he sees it as genetic determinism. In his writing, Lewontin suggests a more nuanced view of evolution is needed, which requires a more careful understanding of the context of the whole organism as well as the environment.
Such concerns about what he views as the oversimplification of genetics has led Lewontin to be a frequent participant in debates, and an active life as a public intellectual. He has lectured widely to promote his views on evolutionary biology and science. In books such as Not in Our Genes (co-authored with Steven Rose and Leon J. Kamin) and numerous articles, Lewontin has questioned much of the claimed heritability of human behavioral traits, such as intelligence as measured by IQ tests.
Some academics have criticized him for rejecting sociobiology for non-scientific reasons. Edward Wilson (1995) suggested that Lewontin's political beliefs affected his scientific view. Lewontin has at times identified himself as Marxist, and admitted that his ideological views have affected his scientific work (Levins and Lewontin 1985). Others such as Kitcher (1985) have countered that Lewontin's criticisms of sociobiology are genuine scientific concerns about the discipline. He wrote that attacking Lewontin's motives amounts to an ad hominem argument. Robert Trivers described Lewontin as "...a man with great talents who often wasted them on foolishness, on preening and showing off, on shallow political thinking and on useless philosophical rumination while limiting his genetic work by assumptions congenial to his politics."
Lewontin has written on the economics of agribusiness. He has contended that hybrid corn was developed and propagated not because of its superior quality, but because it allowed agribusiness corporations to force farmers to buy new seed each year rather than plant seed produced by their previous crop of corn. Lewontin testified in an unsuccessful suit in California challenging the state's financing of research to develop automatic tomato pickers. This favored the profits of agribusiness over the employment of farm workers.
As of 2003, Lewontin was the Alexander Agassiz Research Professor at Harvard. He has worked with and had great influence on many philosophers of biology, including William C. Wimsatt, Elliott Sober, Philip Kitcher, Elisabeth Lloyd, Peter Godfrey-Smith, Sahotra Sarkar, and Robert Brandon, often inviting them to work in his lab. As of mid-2015, Lewontin and his wife Mary Ann live on a farm in Brattleboro, Vermont.
- 1961: Fulbright Fellowship
- 1961: National Science Foundation Senior Postdoctoral Fellow
- 1970s: Membership of the National Academy of Sciences (later resigned)
- 1994: Sewall Wright Award from the American Society of Naturalists
- 2015: Crafoord Prize from the Royal Swedish Academy of Sciences (shared with Tomoko Ohta)
- Lewontin, R. C.; Kojima, K. (Dec 1960). "The Evolutionary Dynamics of Complex Polymorphisms". Evolution (Society for the Study of Evolution) 14 (4): 458–472. doi:10.2307/2405995. JSTOR 2405995.
- Richard C. Lewontin (Jan 1966). "Is Nature Probable or Capricious?". BioScience (University of California Press) 16 (1, Logic in Biological Investigation): 25–27. doi:10.2307/1293548. JSTOR 1293548.
- Lewontin, R. C. (1970). "The Units of Selection". Annual Review of Ecology and Systematics 1: 1–0. doi:10.1146/annurev.es.01.110170.000245.
- "The Apportionment of Human Diversity," Evolutionary Biology, vol. 6 (1972) pp. 391–398.
- Lewontin, R. C. (1974). The genetic basis of evolutionary change. New York: Columbia University Press. ISBN 0-231-03392-3.
- "Adattamento," Enciclopedia Einaudi, (1977) vol. 1, 198–214.
- "Adaptation," Scientific American, vol. 239, (1978) 212–228.
- Gould, S. J.; Lewontin, R. C. (1979). "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme". Proceedings of the Royal Society B: Biological Sciences 205 (1161): 581–98. doi:10.1098/rspb.1979.0086. PMID 42062.
- Lewontin, R. C. (1995). Human diversity (2nd ed.). New York: Scientific American Library. ISBN 0-7167-6013-4.
- "The Organism as Subject and Object of Evolution," Scientia vol. 188 (1983) 65–82.
- Not in Our Genes: Biology, Ideology and Human Nature (with Steven Rose and Leon J. Kamin) (1984) ISBN 0-394-72888-2
- The Dialectical Biologist (with Richard Levins), Harvard University Press (1985) ISBN 0-674-20283-X
- Biology as Ideology: The Doctrine of DNA (1991) ISBN 0-06-097519-9
- The Triple Helix: Gene, Organism, and Environment, Harvard University Press (2000) ISBN 0-674-00159-1
- It Ain't Necessarily So: The Dream of the Human Genome and Other Illusions, New York Review of Books (2000)
- Biology Under The Influence: Dialectical Essays on the Coevolution of Nature and Society (with Richard Levins), (2007)
- Richard Lewontin at the Mathematics Genealogy Project
- Kreitman, Martin Edward (1983). Nucleotide Sequence Variation of Alcohol dehydrogenase in Drosophila melanogaster (PhD thesis). Harvard University.
- Lewontin, R. C.; Hubby, J. L. (1966). "A molecular approach to the study of genic heterozygosity in natural populations. II. Amount of variation and degree of heterozygosity in natural populations of Drosophila pseudoobscura". Genetics 54 (2): 595–609. PMC 1211186. PMID 5968643.
- Hubby, J. L.; Lewontin, R. C. (1966). "A molecular approach to the study of genic heterozygosity in natural populations. I. The number of alleles at different loci in Drosophila pseudoobscura". Genetics 54 (2): 577–594. PMC 1211185. PMID 5968642.
- Lewontin, Richard Charles (2012). The Effects of Population Density and Composition on Viability in Drosophila melanogaster (PhD thesis). Columbia University.
- Lewontin, R. C. and K. Kojima (1960). "The evolutionary dynamics of complex polymorphisms". Evolution 14: 458-472.
- Lewontin, R. C. (1964). "The interaction of selection and linkage. I. General considerations; heterotic models". Genetics 49: 49-67.
- Harris, H. (1966). "Enzyme Polymorphisms in Man". Proceedings of the Royal Society B: Biological Sciences 164 (995): 298–226. doi:10.1098/rspb.1966.0032.
- Kreitman, M. (1983). "Nucleotide polymorphism at the alcohol dehydrogenase locus of Drosophila melanogaster". Nature 304 (5925): 412–417. doi:10.1038/304412a0. PMID 6410283.
- Lewontin, R (1972). "The Apportionment of Human Diversity". Evolutionary Biology 6: 391–398. doi:10.1007/978-1-4684-9063-3_14.
- Edwards, A. W. F. (2003). "Human genetic diversity: Lewontin's fallacy". BioEssays 25 (8): 798–801. doi:10.1002/bies.10315. PMID 12879450.
- Elizabeth Allen et al., 1975, "Against 'Sociobiology'", The New York Review of Books, 13 November 1975
- Gould SJ, Lewontin RC (1979). "The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme". Proc. R. Soc. Lond., B, Biol. Sci. 205 (1161): 581–98. doi:10.1098/rspb.1979.0086. PMID 42062.
- Lewontin RC (1966). "Is nature probable or capricious". Bioscience 16 (1): 25–27. doi:10.2307/1293548.
- Niche Construction: The Neglected Process in Evolution Odling-Smee F.J., Laland K.N., Feldman M.W. Princeton University Press, 2003
- Richard Levins and Richard Lewontin, The Dialectical Biologist, 1985
- Lewontin, R. C. (1978). "Adaptation". Scientific American 239 (3): 212–218, 220, 218 passim. doi:10.1038/scientificamerican0978-212. PMID 705323.
- "Science Contra Darwin", Newsweek, 8 April 1985, p.80
- "Scientist as Activist". http://globetrotter.berkeley.edu/people3/Lewontin/lewontin-con2.html.
- Philip Kitcher (1985). Vaulting Ambition : Sociobiology and the Quest for Human Nature. MIT Press. ISBN 0-262-11109-8.
- Steven Pinker (2002). The Blank Slate: The Denial of Human Nature in Modern Intellectual Life. Penguin Press.
- Rama S. Singh, Costas Krimbas, Diane Paul, and John Beatty (2001). Thinking about Evolution. Cambridge University Press. - a two volume Festschrift for Lewontin with a full bibliography
- Edward O. Wilson (1995). "Science and ideology". Academic Questions 8.
- Richard Lewontin (2004). "The Triple Helix: Gene, Organism and Environment". MIT Press.
- Quotations related to Richard Lewontin at Wikiquote
- an interview given at Berkeley in 2003
- Richard Lewontin's Profile at the California Institute of Technology
- Gene, Organism and Environment: Bad Metaphors and Good Biology - RealAudio stream of Hitchcock lecture on UCTV
- The Concept of Race: The Confusion of Social and Biological Reality - RealAudio stream of Hitchcock lecture on UCTV
- Internalism and Externalism in Biology, lecture delivered at Harvard university on December 13, 2007.