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Richard Charles Lewontin (March 29, 1929 – July 4, 2021) was an American evolutionary biologist, mathematician,
geneticist A geneticist is a biologist or physician who studies genetics, the science of genes, heredity, and variation of organisms. A geneticist can be employed as a scientist or a lecturer. Geneticists may perform general research on genetic process ...
, and social commentator. A leader in developing the
mathematical Mathematics is a field of study that discovers and organizes methods, Mathematical theory, theories and theorems that are developed and Mathematical proof, proved for the needs of empirical sciences and mathematics itself. There are many ar ...
basis of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
and evolutionary theory, he applied techniques from
molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
, such as
gel electrophoresis Gel electrophoresis is an electrophoresis method for separation and analysis of biomacromolecules (DNA, RNA, proteins, etc.) and their fragments, based on their size and charge through a gel. It is used in clinical chemistry to separate ...
, to questions of
genetic variation Genetic variation is the difference in DNA among individuals or the differences between populations among the same species. The multiple sources of genetic variation include mutation and genetic recombination. Mutations are the ultimate sources ...
and evolution. In a pair of seminal 1966 papers co-authored with J. L. Hubby in the journal ''
Genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
'', Lewontin helped set the stage for the modern field of
molecular evolution Molecular evolution describes how Heredity, inherited DNA and/or RNA change over evolutionary time, and the consequences of this for proteins and other components of Cell (biology), cells and organisms. Molecular evolution is the basis of phylogen ...
. In 1979, he and
Stephen Jay Gould Stephen Jay Gould ( ; September 10, 1941 – May 20, 2002) was an American Paleontology, paleontologist, Evolutionary biology, evolutionary biologist, and History of science, historian of science. He was one of the most influential and widely re ...
introduced the term "
spandrel A spandrel is a roughly triangular space, usually found in pairs, between the top of an arch and a rectangular frame, between the tops of two adjacent arches, or one of the four spaces between a circle within a square. They are frequently fil ...
" into
evolutionary theory Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certai ...
. From 1973 to 1998, he held an endowed chair in zoology and biology at Harvard University, and from 2003 until his death in 2021 he was a research professor there. From a sociological perspective, Lewontin strongly opposed genetic determinism and neodarwinism as expressed in the fields of sociobiology and evolutionary psychology. Previously, as a member of Science for the People, he denounced the involvement of prominent scientists in Pentagon programs aimed at developing weapons for the Vietnam War. From the 1990s, he condemned the lobbying of GMOs by the "genetic–industrial complex".


Early life and education

Lewontin was born in New York City to parents descended from late 19th-century
Ashkenazi Jewish Ashkenazi Jews ( ; also known as Ashkenazic Jews or Ashkenazim) form a distinct subgroup of the Jewish diaspora, that Ethnogenesis, emerged in the Holy Roman Empire around the end of the first millennium Common era, CE. They traditionally spe ...
immigrants. His father was a broker of textiles, and his mother a homemaker. He attended Forest Hills High School and the École Libre des Hautes Études in New York. In 1951 he graduated from
Harvard College Harvard College is the undergraduate education, undergraduate college of Harvard University, a Private university, private Ivy League research university in Cambridge, Massachusetts, United States. Part of the Harvard Faculty of Arts and Scienc ...
with a BS degree in
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
. In 1952, Lewontin received an MS degree in mathematical statistics, followed by a
PhD A Doctor of Philosophy (PhD, DPhil; or ) is a terminal degree that usually denotes the highest level of academic achievement in a given discipline and is awarded following a course of graduate study and original research. The name of the deg ...
degree in zoology in 1954, both from
Columbia University Columbia University in the City of New York, commonly referred to as Columbia University, is a Private university, private Ivy League research university in New York City. Established in 1754 as King's College on the grounds of Trinity Churc ...
, where he was a student of
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (; ; January 25, 1900 – December 18, 1975) was a Russian-born American geneticist and evolutionary biologist. He was a central figure in the field of evolutionary biology for his work in shaping the modern ...
. He held faculty positions at
North Carolina State University North Carolina State University (NC State, North Carolina State, NC State University, or NCSU) is a public university, public Land-grant university, land-grant research university in Raleigh, North Carolina, United States. Founded in 1887 and p ...
, the
University of Rochester The University of Rochester is a private university, private research university in Rochester, New York, United States. It was founded in 1850 and moved into its current campus, next to the Genesee River in 1930. With approximately 30,000 full ...
, and the
University of Chicago The University of Chicago (UChicago, Chicago, or UChi) is a Private university, private research university in Chicago, Illinois, United States. Its main campus is in the Hyde Park, Chicago, Hyde Park neighborhood on Chicago's South Side, Chic ...
. In 1973 Lewontin was appointed as Alexander Agassiz Professor of Zoology and Professor of Biology at
Harvard University Harvard University is a Private university, private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the History of the Puritans in North America, Puritan clergyma ...
, holding the position until 1998.


Career


Work in population genetics

Lewontin worked in both theoretical and experimental
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
. A hallmark of his work was an interest in new technology. In 1960, he and Ken-Ichi Kojima gave the equations for change of
haplotype A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
frequencies with interacting natural selection at two loci. 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 Linkage disequilibrium, often abbreviated to LD, is a term in population genetics referring to the association of genes, usually linked genes, in a population. It has become an important tool in medical genetics and other fields In defining LD, it ...
. In 1966, he and J. L. Hubby published a paper that studied the amount of heterozygosity in a population. They used protein
gel electrophoresis Gel electrophoresis is an electrophoresis method for separation and analysis of biomacromolecules (DNA, RNA, proteins, etc.) and their fragments, based on their size and charge through a gel. It is used in clinical chemistry to separate ...
to survey dozens of loci in the fruit fly ''
Drosophila pseudoobscura ''Drosophila pseudoobscura'' is a species of Drosophilidae, fruit fly, used extensively in lab studies of speciation. It is native to western North America. In 2005, ''D. pseudoobscura'' was the second ''Drosophila'' species to have its genome ...
'', 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 Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
. (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. 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 published in 1972, Lewontin identified that most of the variation (80–85%) within human populations is found within local geographic groups, and differences attributable to the " race" groups defined in his study are a minor part of human genetic variability (1–15%). In a 2003 paper,
A. W. F. Edwards Anthony William Fairbank Edwards, Fellow of the Royal Society, FRS One or more of the preceding sentences incorporates text from the royalsociety.org website where: (born 1935) is a British statistician, geneticist and evolutionary biologist. Ed ...
criticized Lewontin's conclusion that race is an invalid taxonomic construct, terming it Lewontin's fallacy. He showed that the probability of racial misclassification of an individual based on variation in a single genetic locus is approximately 30%, but the misclassification probability becomes close to zero if enough loci are studied. That is, it appears that a majority of genetic variation is found within groups only if a single locus is used, but the reverse is true if analyzing a multiplicity of loci. Edwards' paper was commented on by 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."


Affiliations

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 Elliott R. Sober (born 6 June 1948) is an American philosopher. He is noted for his work in philosophy of biology and general philosophy of science. Sober is Hans Reichenbach Professor and William F. Vilas Research Professor Emeritus in the Depar ...
, 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 The National Center for Science Education (NCSE) is a Nonprofit organization, not-for-profit membership organization in the United States whose stated mission is to educate the press and the public on the scientific and educational aspects of con ...
.


Debates within mainstream evolutionary biology

In 1975, when E. O. Wilson's book ''
Sociobiology Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of ...
'' proposed evolutionary explanations for human social behaviors, biologists including Lewontin, his Harvard colleagues
Stephen Jay Gould Stephen Jay Gould ( ; September 10, 1941 – May 20, 2002) was an American Paleontology, paleontologist, Evolutionary biology, evolutionary biologist, and History of science, historian of science. He was one of the most influential and widely re ...
and Ruth Hubbard responded negatively.
Robert Trivers Robert Ludlow "Bob" Trivers (; born February 19, 1943) is an American evolutionary biologist and sociobiologist. Trivers proposed the theories of reciprocal altruism (1971), parental investment (1972), facultative sex ratio determination (197 ...
called these accusations "coming from eminent biologists" "intellectually weak and lazy". Lewontin and Gould introduced the term
spandrel A spandrel is a roughly triangular space, usually found in pairs, between the top of an arch and a rectangular frame, between the tops of two adjacent arches, or one of the four spaces between a circle within a square. They are frequently fil ...
to evolutionary biology, inspired by the architectural term "
spandrel A spandrel is a roughly triangular space, usually found in pairs, between the top of an arch and a rectangular frame, between the tops of two adjacent arches, or one of the four spaces between a circle within a square. They are frequently fil ...
", 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 An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
that exist as a necessary consequence of other (perhaps adaptive) features, but do not directly improve fitness (and thus are not necessarily adaptive). 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 Richard Levins (June 1, 1930 – January 19, 2016) was a Marxist biologist, a population geneticist, biomathematician, mathematical ecologist, and philosopher of science who researched genetic diversity, diversity in human populations. Until his ...
at the University of Chicago), Robert Brandon and Elisabeth Lloyd (who studied with Lewontin as graduate students), Philip Kitcher,
Elliott Sober Elliott R. Sober (born 6 June 1948) is an American philosopher. He is noted for his work in philosophy of biology and general philosophy of science. Sober is Hans Reichenbach Professor and William F. Vilas Research Professor Emeritus in the Depar ...
, 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 ''Darwinism'' is a term used to describe a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others. The theory states that all species of organisms arise and develop through the natural sel ...
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
dialectic Dialectic (; ), also known as the dialectical method, refers originally to dialogue between people holding different points of view about a subject but wishing to arrive at the truth through reasoned argument. Dialectic resembles debate, but the ...
al. M. W. Feldman and others have developed Lewontin's conception in more detailed models under the term
niche construction Niche construction is the ecological process by which an organism alters its own (or another species') local environment. These alterations can be a physical change to the organism’s environment, or it can encompass the active movement of an or ...
. 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 criticized traditional neo-Darwinian approaches to
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
. In his article "Adaptation" in the Italian '' Enciclopedia Einaudi'', and in a modified version for ''
Scientific American ''Scientific American'', informally abbreviated ''SciAm'' or sometimes ''SA'', is an American popular science magazine. Many scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it, with more than 150 Nobel Pri ...
,'' 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 said that his more general, technical criticism of
adaptationism Adaptationism is a scientific perspective on evolution that focuses on accounting for the products of evolution as collections of adaptive traits, each a product of natural selection with some adaptive rationale. A formal alternative would be to ...
grew out of his recognition that the fallacies of
sociobiology Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of ...
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
giraffe The giraffe is a large Fauna of Africa, African even-toed ungulate, hoofed mammal belonging to the genus ''Giraffa.'' It is the Largest mammals#Even-toed Ungulates (Artiodactyla), tallest living terrestrial animal and the largest ruminant on ...
s.


Sociobiology and evolutionary psychology

Along with others, such as Gould, Lewontin was a persistent critic of some themes in neo-Darwinism. Specifically, he criticized proponents of
sociobiology Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of ...
and
evolutionary psychology Evolutionary psychology is a theoretical approach in psychology that examines cognition and behavior from a modern evolutionary perspective. It seeks to identify human psychological adaptations with regard to the ancestral problems they evolved ...
, such as Edward O. Wilson and
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biology, evolutionary biologist, zoologist, science communicator and author. He is an Oxford fellow, emeritus fellow of New College, Oxford, and was Simonyi Professor for the Publ ...
, 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 viewed as the oversimplification of genetics led Lewontin to be a frequent participant in debates, and an active life as a public intellectual. He lectured widely to promote his views on evolutionary biology and science. In the book '' Not in Our Genes'' (co-authored with Steven Rose and Leon J. Kamin) and numerous articles, Lewontin questioned much of the claimed
heritability Heritability is a statistic used in the fields of Animal husbandry, breeding and genetics that estimates the degree of ''variation'' in a phenotypic trait in a population that is due to genetic variation between individuals in that population. T ...
of human behavioral traits, such as
intelligence Intelligence has been defined in many ways: the capacity for abstraction, logic, understanding, self-awareness, learning, emotional knowledge, reasoning, planning, creativity, critical thinking, and problem-solving. It can be described as t ...
as measured by IQ tests. Some academics have criticized him for rejecting
sociobiology Sociobiology is a field of biology that aims to explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within the study of ...
for non-scientific reasons. Edward Wilson (1995) suggested that Lewontin's political beliefs affected his scientific view. Others, such as Kitcher (1985), 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 , short for , refers to several types of arguments that are usually fallacious. Often currently this term refers to a rhetorical strategy where the speaker attacks the character, motive, or some other attribute of the person making an argument ...
'' argument. Lewontin at times identified himself as
Marxist Marxism is a political philosophy and method of socioeconomic analysis. It uses a dialectical and materialist interpretation of historical development, better known as historical materialism, to analyse class relations, social conflic ...
, and asserted that his philosophical views have bolstered his scientific work ( Levins and Lewontin 1985).


Agribusiness

Lewontin has written on the economics of
agribusiness Agribusiness is the industry, enterprises, and the field of study of value chains in agriculture and in the bio-economy, in which case it is also called bio-business or bio-enterprise. The primary goal of agribusiness is to maximize profit ...
. He has contended that hybrid
corn Maize (; ''Zea mays''), also known as corn in North American English, is a tall stout Poaceae, grass that produces cereal grain. It was domesticated by indigenous peoples of Mexico, indigenous peoples in southern Mexico about 9,000 years ago ...
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).


Personal life

As of mid-2015, Lewontin and his wife Mary Jane (Christianson) lived in a log cabin he built with his family in the 1970's in
Marlboro, Vermont Marlboro is a New England town, town in Windham County, Vermont, Windham County, Vermont, United States. The population was 1,722 at the 2020 United States Census, 2020 census. The town is home to the Southern Vermont Natural History Museum and P ...
. Lewontin served many years as a volunteer fireman in Marlboro and also served as a trustee for the Marlboro Music Festival. Dick and Mary Jane had four sons. He was an atheist. Lewontin died at his home in
Cambridge, Massachusetts Cambridge ( ) is a city in Middlesex County, Massachusetts, United States. It is a suburb in the Greater Boston metropolitan area, located directly across the Charles River from Boston. The city's population as of the 2020 United States census, ...
on July 4, 2021, at the age of 92.


Recognition

*1961:
Fulbright Fellowship The Fulbright Program, including the Fulbright–Hays Program, is one of several United States cultural exchange programs with the goal of improving intercultural relations, cultural diplomacy, and intercultural competence between the people o ...
*1961:
National Science Foundation The U.S. National Science Foundation (NSF) is an Independent agencies of the United States government#Examples of independent agencies, independent agency of the Federal government of the United States, United States federal government that su ...
Senior Postdoctoral Fellow *1968: Elected member of the
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, NGO, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the ...
(resigned in 1972) *1994:
Sewall Wright Sewall Green Wright ForMemRS HonFRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongside ...
Award from the American Society of Naturalists *2015:
Crafoord Prize The Crafoord Prize () is an annual science prize established in 1980 by Holger Crafoord, a Swedish industrialist, and his wife Anna-Greta Crafoord following a donation to the Royal Swedish Academy of Sciences. It is awarded jointly by the Acade ...
from the
Royal Swedish Academy of Sciences The Royal Swedish Academy of Sciences () is one of the Swedish Royal Academies, royal academies of Sweden. Founded on 2 June 1739, it is an independent, non-governmental scientific organization that takes special responsibility for promoting nat ...
(shared with Tomoko Ohta) *2017: Thomas Hunt Morgan Medal from the Genetics Society of America


Bibliography

* * * * " The Apportionment of Human Diversity," ''Evolutionary Biology'', vol. 6 (1972) pp. 391–398. * * "Adattamento," ''Enciclopedia Einaudi'', (1977) vol. 1, 198–214. * "Adaptation," ''Scientific American'', vol. 239, (1978) 212–228. * * * "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) . *'' The Dialectical Biologist'' (with
Richard Levins Richard Levins (June 1, 1930 – January 19, 2016) was a Marxist biologist, a population geneticist, biomathematician, mathematical ecologist, and philosopher of science who researched genetic diversity, diversity in human populations. Until his ...
), Harvard University Press (1985) . *''Biology as Ideology: The Doctrine of DNA'' (1991) . *''The Triple Helix: Gene, Organism, and Environment'', Harvard University Press (2000) . *''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 Richard Levins (June 1, 1930 – January 19, 2016) was a Marxist biologist, a population geneticist, biomathematician, mathematical ecologist, and philosopher of science who researched genetic diversity, diversity in human populations. Until his ...
), (2007).
''Agricultural research and the penetration of capital''
(1982), '' Science for the People'' 14 (1): 12–17 * Lewontin, R.C. ''The maturing of capitalist agriculture: farmer as proletarian''. P. 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.


References


Further reading

* * - a two volume Festschrift for Lewontin with a full bibliography * *


External links

*
an interview given at Berkeley in 2003


at the
California Institute of Technology The California Institute of Technology (branded as Caltech) is a private research university in Pasadena, California, United States. The university is responsible for many modern scientific advancements and is among a small group of institutes ...

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. * {{DEFAULTSORT:Lewontin, Richard 1929 births 2021 deaths 20th-century American mathematicians 21st-century American mathematicians 21st-century American biologists American atheists American Marxists Columbia University faculty Columbia University alumni American critics of creationism American evolutionary biologists Extended evolutionary synthesis Harvard College alumni Harvard University faculty American intelligence researchers Jewish American atheists Jewish American scientists North Carolina State University faculty People from Brattleboro, Vermont Population geneticists People involved in race and intelligence controversies Santa Fe Institute people Scientists from New York City Scientists from Vermont American theoretical biologists University of Chicago faculty University of Rochester faculty Mathematicians from New York (state) Presidents of the American Society of Naturalists