Richard C. Lewontin (1929–2021) : At the Cutting Edge of Biology and the Relationship Between Science and Race

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  • Thanks to Toyin Falola for his Post that Brought Richard  Lewontin to My Attention



  • OBITUARY
  • 13 July 2021

Richard C. Lewontin (1929–2021)

Pioneer of molecular evolution who campaigned against biological racism.
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Black and white photo of Richard C. Lewontin standing at a chalk board

Credit: Ernst Mayr Library and Archives of the Museum of Comparative Zoology, Harvard University

Richard Lewontin was a groundbreaking geneticist, best known for bringing molecular tools into evolutionary biology and for his advocacy against the use of science to rationalize structural inequity. Lewontin and his collaborators revealed how natural selection acts to shape variation, exploring its effect on genes, groups and individuals. Moving between mathematical and statistical analysis, fieldwork and laboratory experiment, they set the course of molecular population genetics. Lewontin saw no place for his discipline in attempts to explain why “the children of oil magnates tend to become bankers, while the children of oil workers tend to be in debt to banks”.

Lewontin’s sometimes controversial critiques of science, often from a Marxist perspective, inspired new thinking on the relationship between science, politics and society. He was an outspoken critic of sociobiology and adaptationism (the idea that all traits evolved as adaptations of an organism to its environment). He despised the use of biology to justify racist ideology, especially with regard to IQ testing. His celebrated essay ‘The spandrels of San Marco and the Panglossian paradigm’, written with his colleague Stephen Jay Gould (Proc. R. Soc. Lond. B. 205, 581–598; 1979) skewered, among other things, a “reliance upon plausibility alone as a criterion for accepting speculative tales”. Lewontin has died aged 92.

Richard Lewontin was born into an upper-middle-class Jewish family in New York City, and originally studied biology at Harvard University in Cambridge, Massachusetts, in the early 1950s. At the time, Harvard had no faculty member specializing in genetics, so Lewontin studied with a visitor, Leslie C. Dunn, from Columbia University in New York City. Dunn persuaded Lewontin to join the Columbia laboratory of Theodosius Dobzhansky, then the most influential evolutionary geneticist in the world. Lewontin adopted Dobzhansky’s investigation of the nature of selection and its impact on the variability of natural and laboratory populations. He completed his PhD in 1954.

That year, Lewontin joined the faculty at North Carolina State University in Raleigh. Here, he focused primarily on mathematical population genetics and worked with Ken-Ichi Kojima on genetic linkage, the tendency of neighbouring genetic sequences to be inherited together. After periods at the University of Rochester, New York, and the University of Chicago, Illinois, he spent the rest of his career at Harvard.

During his time at Rochester in the early 1960s, attempts to study genetic variation in natural populations were approaching an impasse. On a visit to the University of Chicago, Lewontin met Jack Hubby, who was adapting the biochemical technique of electrophoresis (which separates molecules by charge and size) to study the fruit fly Drosophila. They realized that detecting small differences between proteins could provide a new means of measuring genetic variability.

Lewontin moved to the University of Chicago and, with Hubby, published two landmark papers (Genetics 54, 577–594 and 595–609; 1966), which opened the way for the widespread application of electrophoresis and marked the beginning of molecular population genetics. These papers also revealed higher than expected amounts of genetic variability, addressing a long-standing dispute about whether natural selection maintains genetic variability in natural populations. In 1984, Martin Kreitman, working between Lewontin’s and Walter Gilbert’s laboratories at Harvard, brought DNA sequencing to bear on this question.

In Chicago in the 1960s, Lewontin became increasingly politically active, speaking out against racial discrimination, the Vietnam War and economic inequality. His fervent convictions led him to renounce his election to the US National Academy of Sciences, because of its support for secret war research. With ecologist Dick Levins and support from the Ford Foundation, he assembled a group to investigate the role of capital in agricultural research, such as the development of hybrid crop plants. Lewontin and Levins’s collaboration also led to a series of essays on biology and society from a Marxist perspective, published later as The Dialectical Biologist (1985) and Biology Under the Influence (2007). Like his critiques of sociobiology, many of these essays treated science as politics, arguing against reductionism and determinism that favoured biological explanations of complex biosocial phenomena.

Lewontin also spoke up against biological racism. His landmark paper ‘The Apportionment of Human Diversity’ (in Evolutionary Biology Vol. 6 (eds T. Dobzhansky et al.) Springer, 1972) found more variation within so-called ‘racial groups’ than between them, leading him to argue that such distinctions had no genetic basis. When biological arguments for race were again put forward in the context of mental testing in the 1980s, he opposed them on scientific and social grounds, notably in Not in Our Genes: Biology, Ideology, and Human Nature (1984), co-authored with Steven Rose and Leon Kamin, and reissued in 2017 during the administration of US President Donald Trump. He continued to publish in this realm for decades.

Lewontin described himself as a pessimistic biologist. He was a profoundly critical thinker, willing to challenge the scientific and philosophical foundations of his discipline as well as their social, cultural and political consequences. His research and reflections set an agenda for generations of biologists, philosophers of biology and socially engaged scholars.

In keeping with his socialism, he disliked biography and its celebration of the individual. When, in 1997, I asked him how I should write about his life, he pulled out of his desk a list of every graduate student, postdoc and visitor at his laboratory — more than 100 people — and said I should write about all of them. They were his greatest source of pride as a scientist.

Nature 595, 489 (2021)

doi: https://doi.org/10.1038/d41586-021-01936-6

Richard Lewontin

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Richard Lewontin
Born
Richard Charles Lewontin

March 29, 1929
DiedJuly 4, 2021 (aged 92)
Alma materHarvard University (BS)
Columbia University (PhD)
Known forEvolutionary biology
Population genetics
AwardsSewall Wright Award (1994), Crafoord Prize (2015), Thomas Hunt Morgan Medal (2017)
Scientific career
FieldsGenetics
Evolutionary biology
Population genetics
InstitutionsHarvard University
North Carolina State University
University of Rochester
University of Chicago
Columbia University
ThesisThe Effects of Population Density and Composition on Viability in Drosophila melanogaster (1955)
Doctoral advisorTheodosius Dobzhansky[1]
Doctoral studentsAdriana Briscoe
Jerry Coyne
Joseph Felsenstein
Martin Kreitman[2]
Russell Lande

Richard Charles Lewontin (March 29, 1929 – July 4, 2021[3]) was an American evolutionary biologist, mathematician, geneticist, 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,[4][5] 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 from 2003 until his death in 2021 had been a research professor there.

Lewontin opposed genetic determinism.[6]

Contents

Early life and education[edit]

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 with a BS degree in biology. In 1952, Lewontin received an MS degree in mathematical statistics, followed by a PhD degree in zoology in 1954,[7] 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.

Career[edit]

Work in population genetics[edit]

Lewontin worked in both theoretical and experimental population genetics. A hallmark of his work was an interest in new technology. He was the first person to do a computer simulation of the behavior of a single gene locus (previous simulation work having been of models with multiple loci).[citation needed] 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.[8] 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.[9] (He also introduced the term "linkage disequilibrium", about which many population geneticists have been unenthusiastic.[10])

In 1966, he and J. L. Hubby published a paper that revolutionized population genetics.[4] They used protein gel electrophoresis to survey dozens of loci in the fruit fly 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.)[11] 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.[12]

Work on human genetic diversity[edit]

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%).[13] 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.[14] Edwards' criticism in turn garnered its own criticism from biologists such as Jonathan Marks, who argued that "the point of the theory of race was to discover large clusters of people that are principally homogeneous within and heterogeneous between, contrasting groups. Lewontin's analysis shows that such groups do not exist in the human species, and Edwards' critique does not contradict that interpretation."[15]

Critique of mainstream evolutionary biology[edit]

In 1975, when E. O. Wilson's book Sociobiology proposed evolutionary explanations for human social behaviors, biologists including Lewontin, his Harvard colleagues Stephen Jay Gould and Ruth Hubbard responded negatively.[16]

Lewontin and 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).[17] 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?".[18]

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 and others[19] have developed Lewontin's conception in more detailed models under the term niche construction.

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.[20]

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 modified version for 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.[21] 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.

Lewontin accused neo-Darwinists of telling Just-So Stories when they try to show how natural selection explains such novelties as long-necked giraffes.[22]

Sociobiology and evolutionary psychology[edit]

Along with others, such as Gould, Lewontin has been a persistent critic of some themes in neo-Darwinism. Specifically, he has criticized 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.[23]

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.[citation needed]

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. 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."[24] 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.[citation needed] Lewontin has at times identified himself as Marxist, and asserted that his philosophical views have bolstered his scientific work (Levins and Lewontin 1985).

Agribusiness[edit]

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 1982). 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 (Lewontin 2000).

Lewontin, R. C. 1982. Agricultural research and the penetration of capital. Science for the People 14 (1): 12–17. http://www.science-for-the-people.org/wp-content/uploads/2015/07/SftPv14n1s.pdf.

Lewontin, R.C. 2000. The maturing of capitalist agriculture: farmer as proletarian. Pgs 93–106 in F. Magdoff, J. B. Foster, and F. H. Buttel, Eds. 2000. Hungry for Profit: The Agribusiness Threat to Farmers, Food, and the Environment. Monthly Review Press, NY.

Personal life[edit]

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.

Since 2013, Lewontin has been listed on the Advisory Council of the National Center for Science Education.[25]

As of mid-2015, Lewontin and his wife Mary Jane lived on a farm in Brattleboro, Vermont. He was an atheist.[26]

Lewontin died on July 4, 2021, at the age of 92.[3][27]

Recognition[edit]

Bibliography[edit]

Scholia has a profile for Richard Lewontin (Q659265).

References[edit]

  1. ^ Richard Lewontin at the Mathematics Genealogy Project
  2. ^ Kreitman, Martin Edward (1983). Nucleotide Sequence Variation of Alcohol dehydrogenase in Drosophila melanogaster (PhD thesis). Harvard University. ProQuest 303271509.
  3. ^ Jump up to:a b Angier, Natalie (July 7, 2021). "Richard C. Lewontin, Eminent Geneticist With a Sharp Pen, Dies at 92". The New York Times. Retrieved July 8, 2021.
  4. ^ Jump up to:a b 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. doi:10.1093/genetics/54.2.595. PMC 1211186. PMID 5968643.
  5. ^ 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. doi:10.1093/genetics/54.2.577. PMC 1211185. PMID 5968642.
  6. ^ Peters, Ted (2003). Playing God? genetic determinism and human freedom (2nd ed.). New York: Routledge. pp. 29–31. ISBN 978-0-415-94249-2.
  7. ^ Lewontin, Richard Charles (2012). The Effects of Population Density and Composition on Viability in Drosophila melanogaster (PhD thesis). Columbia University. ProQuest 301991815.
  8. ^ Lewontin, R. C.; Kojima, Kenichi (1960). "The evolutionary dynamics of complex polymorphisms". Evolution 14: 458-472.
  9. ^ Lewontin, R. C. (1964). "The interaction of selection and linkage. I. General considerations; heterotic models". Genetics 49: 49-67.
  10. ^ Slatkin, Montgomery (June 2008). "Linkage disequilibrium — understanding the evolutionary past and mapping the medical future" (PDF). Nature Reviews Genetics. 9(6): 477–485. doi:10.1038/nrg2361. PMC 5124487. PMID 18427557.
  11. ^ Harris, H. (1966). "Enzyme Polymorphisms in Man". Proceedings of the Royal Society B: Biological Sciences. 164 (995): 298–310. Bibcode:1966RSPSB.164..298H. doi:10.1098/rspb.1966.0032. PMID 4379519. S2CID 43582403.
  12. ^ Kreitman, M. (1983). "Nucleotide polymorphism at the alcohol dehydrogenase locus of Drosophila melanogaster". Nature. 304 (5925): 412–417. Bibcode:1983Natur.304..412K. doi:10.1038/304412a0. PMID 6410283. S2CID 4348580.
  13. ^ Lewontin, R (1972). "The Apportionment of Human Diversity". Evolutionary Biology. 6: 391–398. doi:10.1007/978-1-4684-9063-3_14. ISBN 978-1-4684-9065-7.
  14. ^ Edwards, A. W. F. (2003). "Human genetic diversity: Lewontin's fallacy". BioEssays. 25 (8): 798–801. doi:10.1002/bies.10315. PMID 12879450.
  15. ^ Marks, Jonathan M. (2010). "Ten Facts about Human Variation". In Muehlenbein, M. P. (ed.). Human Evolutionary Biology. Cambridge University Press. p. 270. ISBN 9781139789004.
  16. ^ Elizabeth Allen et al., 1975, "Against 'Sociobiology'", The New York Review of Books, November 13, 1975
  17. ^ 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. Bibcode:1979RSPSB.205..581G. doi:10.1098/rspb.1979.0086. PMID 42062. S2CID 2129408.
  18. ^ Lewontin RC (1966). "Is nature probable or capricious". BioScience. 16 (1): 25–27. doi:10.2307/1293548. JSTOR 1293548.
  19. ^ Niche Construction: The Neglected Process in Evolution Odling-Smee F. J., Laland K. N., Feldman M. W. Princeton University Press, 2003
  20. ^ Richard Levins and Richard Lewontin, The Dialectical Biologist, 1985
  21. ^ Lewontin, R. C. (1978). "Adaptation". Scientific American. 239 (3): 212–218, 220, 218 passim. Bibcode:1978SciAm.239c.212L. doi:10.1038/scientificamerican0978-212. PMID 705323.
  22. ^ "Science Contra Darwin", Newsweek, April 8, 1985, p.80
  23. ^ Lewontin, Richard; Leon Kamin; Steven Rose (1984). Not in Our Genes: Biology, Ideology, and Human Nature. Pantheon Books. ISBN 0-394-50817-3.
  24. ^ [1]
  25. ^ "Advisory Council". ncse.com. National Center for Science Education. Archived from the original on August 10, 2013. Retrieved October 30, 2018.
  26. ^ "The Wars Over Evolution" New York Review of Books October 20, 2005 "I, his student and scientific epigone, ingested my unwavering atheism..."
  27. ^ "Richard Lewontin: Pioneering evolutionary biologist dies aged 92". New Scientist. July 5, 2021. Retrieved July 5, 2021.
  28. ^ "Scientist as Activist". globetrotter.berkeley.edu. Institute of International Studies. Archived from the original on August 11, 2004. Retrieved October 31, 2018.
  29. ^ "Genetics Society of America honors Richard Lewontin with 2017 Thomas Hunt Morgan Medal". eurekalert.org. Eurekalert. Archived from the original on March 24, 2017. Retrieved October 31, 2018.

Further reading[edit]

External links[edit]

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