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Units of Selection and the Structure of the Multi-Level Genome

Published online by Cambridge University Press:  21 March 2022

William C. Wimsatt*
Affiliation:
Department of Philosophy, Committee on Conceptual Foundations of Science and Committee on Evolutionary Biology, University of Chicago

Extract

The dominant view among evolutionary biologists today is that the gene is the only unit of selection. According to this view, larger units are too unstable in evolutionary time to act as units of selection. Chromosomes are broken up by sexual recombination. At the level of the individual organism, phenotypes pass on genes, but (it is claimed!) last themselves only one generation. Genotypes of individuals, collections of chromosomes, are rearranged by Mendelian independent assortment in sexual reproduction and so also persist for only one generation. Groups of individuals are still more ephemeral, or so we are told.

All of these points are argued at length by George C. Williams in his book Adaptation and Natural Selection which, since its publication in 1966, has been the watershed for the rise to dominance of this reductionistic vision of evolutionary theory.

Type
Part II. Unity of Science— Group Selection and Sociobiology
Copyright
Copyright © 1981 by the Philosophy of Science Association

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Footnotes

1

The debts I owe for this paper are similar to those for my earlier paper, on the topic (1980b). Intellectually the viewpoints expressed here owe most to Richard Lewontin and Richard Levins, and more recently to Michael Wade. To James Crow and to Russ Lande, I owe special debts of thanks, for each prevented me Cat places indicated in the text) from making major errors. Unfortunately, the errors avoided in this paper (my prior belief that most population geneticists believed that most variance in fitness is additive), is mistakenly attributed to Crow in 1980b, p. 237. I here admit my error, and thank him for his many useful comments on the topic, most of which have been incorporated in the present discussion. Others whose comments on the earlier paper.were very useful to me in writing this one include James Griesemer, Jack Hirshleifer, David Hull, Marcy Lawton, Richard Michod, Bob Richardson, and Elliott Sober. Hull, Sober and I may not be in complete agreement, tut the at least approximate consilience of our views (relative to those of the “reductionistic opposition“) has been a productive spur to further work to reconcile our differences. Finally, the vast majority of this work was done with support from the National Science Foundation, under grant NSF-S0C78-07310. I thank them for this generous support. Frances LaDuke has again beautifully typed this paper.

References

Allison, A.C. (1956). “Population Genetics of Abnormal Human Haemoglobins.” Acta Genetica 6: 430-434.Google ScholarPubMed
Ayala, F.J. (ed.). (1976). Molecular Evolution. Sunderland, Mass.: Sinauer.Google Scholar
Bonner, J.T. (1965). Size and Cycle. Princeton: Princeton University Press.CrossRefGoogle Scholar
Carlson, E.A. (1966). The Gene: A Critical History. Philadelphia: Saunders.Google Scholar
Cassidy, John. (1978). “Philosophical Aspects of the Group Selection Controversy.” Philosophy of Science 45: 575-594.CrossRefGoogle Scholar
Cavalli-Sforza, L.L. and Feldman, M.W. (1981). Cultural Transmission and Evolution: A Quantitative Approach. Princeton: Princeton University Press.Google ScholarPubMed
Crow, James F. and Kimura, Motoo. (1970). An Introduction to Population Genetic Theory. New York: Harper and Row.Google Scholar
Darwin, Charles. (1859). The Origin of Species. London: John Murry. (Faosimilie reprint of the first edition with introduction by Mayr, Ernst, Harvard University Press, 1964.)Google Scholar
Dawkins, R. (1976). The Selfish Gene. New York: Oxford University Press.Google Scholar
Dawkins, R. (1978). “Replicator Selection and the Extended Phenotype.” Zeitschrift für Tierpsychologie 47: 61-76.CrossRefGoogle ScholarPubMed
Dickerson, R.C. (1972). “The History and Structure of an Ancient ProteinScientific American 226(4): 58-72.CrossRefGoogle ScholarPubMed
Falconer, D.S. (1977). “Why Are Mice the Size They Are?” In Proceedings of the International Congress on Quantitative Genetics. Edited by Pollack, E. Kempthorne, O. and Bailey, T.B. Ames, Jr. Iowa: The Iowa State University Press. Pages 19-21.Google Scholar
Fisher, R.A. (1930). The Genetioal Theory of Natural Selection. Oxford: The Clarendon Press.Google Scholar
Gould, Stephen Jay. (1977). Ontogeny and Phylogeny. Cambridge: Harvard University Press.Google Scholar
Hamilton, W.D. (1964a). “The Genetical Evolution of Social Behavior, I.Journal of Theoretical Biology 7: 1-16.CrossRefGoogle Scholar
Hamilton, W.D. (1964b). “The Genetical Evolution of Social Behavior, II.” Journal of Theoretical Biology 7: 17-52.CrossRefGoogle Scholar
Hartl, Daniel L. (1980). Principles of Population Genetics. Stamford, Conn.: Sinauer.Google Scholar
Hull, David L. (1980). “Individuality and Selection.” In Annual Review of Ecology and Systematics. Volume 11. Edited by Johnston, R.F., et al. Pages 311-332.CrossRefGoogle Scholar
Hull, David L. (1981a). “The Herd as a Means.” In PSA 1980, Volume 2. Edited by Asquith, P.D. and Giere, R.N. . East Lansing, Michigan: Philosophy of Science Association. Pages 73-92.Google Scholar
Hull, David L. (1981b). “The Naked Meme.” Working paper for the conference on “Epistemologically Relevant Internalist Sociology of Science,” Syracuse University, June 20-26, 1981.Google Scholar
Jenkin, Fleeming. (1867). “Review of Darwin's Origin of Species.The Worth British Review, new series , 7 (March-June): 277-318. As Reprinted in Darwin and His Critics. Edited by Hull, D. .Cambridge, Mass.: Harvard University Press, 1973. Pages 302-350.Google Scholar
Kempthorne, Oscar. (1969). An Introduction to Genetic Statistics. Ames: Iowa State University Press.Google Scholar
Levene, H., Pavlovsky, O., and Dobzhansky, T. (1951). “Interaction of the Adaptive Values in Polymorphic Experimental Populations of Drosophila Pseudoobscura.” Evolution 8: 335-319.Google Scholar
Levins, Richard. (1966). “The Strategy of Model Building in Population Biology.” American Scientist 54: 421-431.Google Scholar
Levins, Richard.(1968). Evolution in Changing Environments. Princeton: Princeton University Press.CrossRefGoogle Scholar
Levins, Richard.. (1973). “The Limits of Complexity.” In Hierarchy Theory: The Challenge of Complex Systems. Edited by Pattee, Howard H. , New York: George Braziller. Pages 109-127.Google Scholar
Lewontin, Richard C. (1958). “A General Method for Investigating the Equilibrium of Gene Frequency in a Population.” Genetics 43: 419-434.CrossRefGoogle ScholarPubMed
Lewontin, Richard C. (1970). “The Units of Selection.” Annual Review of Ecology and Systematics 1: 1-18.CrossRefGoogle Scholar
Lewontin, Richard C . (1974). The Genetic Basis of Evolutionary Change. New York: Columbia University Press.Google Scholar
Lewontin, Richard C. (1978). “Adaptation.”Scientific American 239 (3) : 212-230.CrossRefGoogle ScholarPubMed
Lumsden, C.J., and Wilson, E.O. (1981). Genes, Mind and Culture: The Revolutionary Process. Cambridge: Harvard University Press.Google Scholar
Masters, R.D. (1970). “Genes, Language, and Evolution.” Semiotica 2: 295-320.CrossRefGoogle Scholar
Maynard Smith, John. (1973). John Mavnard Smith on Evolution. Edinburgh: Edinburgh University Press.Google Scholar
Maynard Smith, John. (1975). The Theory of Evolution. 3rd ed. London: Penguin.Google Scholar
Maynard Smith, John. (1978). The Evolution of Sex. Cambridge: Cambridge University Press.Google Scholar
Mayr, Ernst. (1974). “Behavior Programs and Evolutionary Strategies.” American Scientist 62: 650-659.Google ScholarPubMed
McCauley, D.E., and Wade, M.J. (1980). “Group Selection: The Genetic and Demographic Basis for the Phenotypic Differentiation of Small Populations of Tribolium castaneum.” Evolution 34: 813-821.CrossRefGoogle ScholarPubMed
Michod, Richard. (1981). “The Genetical Theory of Social Behavior.” Annual Review of Ecology and Svstematics. Forthcoming.Google Scholar
Nagylaki, Thomas. (1977). Selection in One and Two-Locus Systems. Lecture Notes in, Biomathematics, Volume 15. Berlin: Springer Verlag.CrossRefGoogle Scholar
Nickles, Thomas (ed.). (1980a). Scientific Discovery. Volume I: General and Methodological Studies. Dordrecht: Reidel.CrossRefGoogle Scholar
Nickles, Thomas. (1980b). Scientific Discovery. Volume II: Historical and Scientific Case Studies. Dordrecht: Reidel.CrossRefGoogle Scholar
Nickles, Thomas. (1981a). “What Is the Problem that We Might Solve It?Synthese 47: 85-118.CrossRefGoogle Scholar
Nickles, Thomas . (1981b). “Heuristic Appraisal, Discovery and Justification: Whither Philosophy of Science?” Forthcoming in Laudan, L. , ed., Pittsburgh Studies in Philosophy of Science.Google Scholar
Provine, William B. (1971). The Origins of Theoretical Population Genetics. Chicago: The University of Chicago Press.Google Scholar
Roughgarden, Jonathan. (1979). Theory of Population Genetics and Evolutionary Ecology: An Introduction. New York: MacMillan.Google Scholar
Salmon, W.C. (1971). Statistical Explanation and Statistical. Relevance. Pittsburgh: University of Pittsburgh Press.CrossRefGoogle Scholar
Simon, Herbert A. (1969). The Sciences of the Artificial. Cambridge, Mass.: Massachusetts Institute of Technology Press.Google Scholar
Sober, Elliott . (1981). “Holism, Individualism and the Units of Selection.” In PSA 1980, Volume 2. Edited by Asquith, P.D and Giere, R.N.. East Lansing, Michigan: Philosophy of Science Association. Pages 93-121.Google Scholar
Throckmorton, Lynn H. (1978). “Molecular Phylogenetics.” Beltsville Symposia in Agricultural Research. 2. Blosystematios in Agriculture. Edited by Romberger, J.A., Foote, R.H., Knutson, L. and Lenta, P.L. Montclair, N.J.: Allanheld, Osmun & Co., and New York: John Wiley & Sons. Pages 221-239.Google Scholar
Wade, Michael J. (1978). “A Critical Review of the Models of Group Selection.” Quarterly Review of Biology 53; 101-114.CrossRefGoogle Scholar
Wade, Michael J. (1979). “The Evolution of Social Interactions by Family Selection.” American Naturalist 113: 399-417.CrossRefGoogle Scholar
Wade, M.J., and McCauley, D.E. (1980). “Group Selection: The Phenotypic and Genotypic Differentiation of Small Populations.” Evolution 34: 799-812.CrossRefGoogle ScholarPubMed
Wade, M.J., and Wimsatt, W.C. (1981). “The Blending Theory of Group Inheritance.” Evolution. Forthcoming.Google Scholar
Wallace, B. (1968). Topics in Population Genetics. New York: Norton.Google Scholar
Williams, George C. (1966). Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton: Princeton University Press.Google Scholar
Wimsatt, William C. (1970). “Review of G.C. Williams’ Adaptation and Natural Selection.” Philosophy of Science 37: 620-623.CrossRefGoogle Scholar
Wimsatt, William C. (1974). “Complexity and Organization.” In PSA-1972 (Boston Studies in The Philosophy of Science. Volume XX). Edited by Schaffner, K.F. and Cohen, R.S.. Dordrecht: Reidel. Pages 67-86.CrossRefGoogle Scholar
Wimsatt, William C.(1976a). “Reductionism, Levels of Organization, and the Mind-Body Problem.” In Consciousness and the Brain: Scientific and Philosophical Strategies. Edited by Globus, G.G., Maxwell, G., and Savodnik, I.. New York: Plenum. Pages 199-267.Google Scholar
Wimsatt, William C. (1976b). “Reductive Explanation: A Functional Account.” In PSA - 1974 (Boston Studies in the Philosophy of Science, Volume XXXII). Edited by Cohen, R.S., Hooker, C.A., Michalos, A.C. and van Evra, J.W.. Dordrecht: Reidel. Pages 671-710.CrossRefGoogle Scholar
Wimsatt, William C. (1979). “Reduction and Reductionism.” In Current Research in Philosophy of Science. Edited by Asquith, P.D. and Kyburg, H. Jr. East Lansing, Michigan: Philosophy of Science Association. Pages 352-377.Google Scholar
Wimsatt, William C.(1980a). “Randomness and Perceived-Randomness in Evolutionary Biology.” Synthese 43: 287-329.CrossRefGoogle Scholar
Wimsatt, William C. (1980b). “Reductionistic Research Strategies and Their Biases in the Units of Selection Controversy.” In Scientific Discovery. Volume II; Historical and Scientific Case Studies. Edited by Nickles, Thomas. 213-259. Dordrecht: Reidel. Pages 213-259.CrossRefGoogle Scholar
Wimsatt, William C. (1981). “Robustness, Reliability and Multiple Determination in Science.” In Scientific Inquiry and the Social Sciences: A Volume in Honor of Donald T. Campbell. Edited by Brewer, M. and Collins, B. . San Francisco: Jossey-Bass. Pages 124-163.Google Scholar
Wright, Sewall, . (1968). Genetic and Blometric Foundations (Evolution and the Genetics of Populations. Volume 1.) Chicago: University of Chicago Press.Google Scholar
Wright, Sewall.(1978). Variability Within And Among Natural Populations (Evolution and the Genetics of Populations, Volume 4.) Chicago: University of Chicago Press.Google Scholar