Biology and Philosophy

, Volume 14, Issue 2, pp 167–180 | Cite as

The Units of Selection Revisited: The Modules of Selection

  • Robert N. Brandon


Richard Lewontin's (1970) early work on the “units” of selection initiated the conceptual and theoretical investigations that have led to the hierarchical perspective on selection that has reached near consensus status today. This paper explores other aspects of his work, work on what he termed “continuity” and “quasi-independence”, that connect to contemporary explorations of modularity in development and evolution. I characterize such modules and argue that they are the true units of selection in that they are what evolution by natural selection individuates, selects among, and transforms.

levels of selection modularity units of selection 


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  1. Altenberg, L.: 1995, 'Genome Growth and the Evolution of the Genotype-phenotype Map', in W. Banzhaf and F. H. Eeckman (eds.), Evolution and Biocomputation: Computational Models of Evolution, Springer-Verlag, Berlin, pp. 205-259.Google Scholar
  2. Bonner, J. T.: 1988, The Evolution of Complexity, Princeton University Press, Princeton.Google Scholar
  3. Brandon, R. N.: 1982, 'The Levels of Selection', in P. Asquith and T. Nickles (eds.), PSA 1982, Vol. 1, Philosophy of Science Association, East Lansing, MI, pp. 315-323.Google Scholar
  4. Brandon, R. N.: 1990, Adaptation and Environment, Princeton University Press, Princeton.Google Scholar
  5. Brandon, R. N.: 1997, 'Discussion: Reply to Hitchcock', Biology and Philosophy 12, 531-538.Google Scholar
  6. Brandon, R. N.: forthcoming, 'Organism and Environment Revisited', in R. Singh, D. Paul, C. Krimbas and J. Beatty (eds.), Thinking about Evolution: Historical, Philosophical, and Political Perspectives, Cambridge University Press, Cambridge.Google Scholar
  7. Brandon, R. N. and Burian, R. M. (eds.): 1984, Genes, Organisms, Populations: Controversies Over the Units of Selection, The MIT Press, Cambridge, MA.Google Scholar
  8. Brandon, R. N., Antonovics, J., Burian, R., Carson, S., Cooper, G., Davies, P. S., Horvath, C., Mishler, B. D., Richardson, R. C., Smith, K. and Thrall, P.: 1994, 'Discussion: Sober on Brandon on Screening-off and the Levels of Selection', Philosophy of Science 61, 475-486.Google Scholar
  9. Brandon, R. N. and Antonovics, J.: 1996, 'The Coevolution of Organism and Environment', in R. N. Brandon, Concepts and Methods in Evolutionary Biology, Cambridge University Press, Cambridge, pp. 161-178.Google Scholar
  10. Darwin, C.: 1871, The Descent of Man, John Murray, London.Google Scholar
  11. Dawkins, R.: 1976, The Selfish Gene, Oxford University Press, Oxford.Google Scholar
  12. Dawkins, R.: 1982, 'Replicators and Vehicles', in King's College Sociobiology Group (eds.), Current Problems in Sociobiology, Cambridge University Press, Cambridge, pp. 45-64.Google Scholar
  13. Dawkins, R.: 1989, 'The Evolution of Evolvability', in C. G. Langton (ed.), Artificial Life, the Proceedings of an Interdisciplinary Workshop on the Synthesis and Simulation of Living Systems, Addison-Wesley, Redwood City, CA, pp.Google Scholar
  14. Holland, J. H.: 1992, Adaptation in Natural and Artificial Systems, MIT Press, Cambridge, MA.Google Scholar
  15. Hull, D.: 1981, 'Units of Evolution: A Metaphysical Essay', in U. L. Jensen and R. Harré (eds.), The Philosophy of Evolution, Harvester Press, Brighton.Google Scholar
  16. Kauffman, S. A.: 1989, 'Adaptation on Rugged Fitness Landscapes', in D. Stein (ed.), Lectures in the Science of Complexity, Vol. 1, Addison-Wesley, Longman, Reading, MA, pp. 527-618.Google Scholar
  17. Lewontin, R. C.: 1970, 'The Units of Selection', Annual Review of Ecology and Systematics 1, 1-18.Google Scholar
  18. Lewontin, R. C.: 1974, The Genetic Basis of Evolutionary Change, Columbia University Press, New York.Google Scholar
  19. Lewontin, R. C.: 1978; 'Adaptation', Scientific American 239, 156-169.Google Scholar
  20. Lewontin, R. C.: 1983a, 'Gene, Organism and Environment', in D. S. Bendall (ed.), Evolution from Molecules to Men, Cambridge University Press, Cambridge, pp. 273-285.Google Scholar
  21. Lewontin, R. C.: 1983b, 'The Organism as the Subject and Object of Evolution', Scientia 118, 63-82.Google Scholar
  22. Lloyd, E. A.: 1998, The Structure and Confirmation of Evolutionary Theory, Greenwood Press, New Your.Google Scholar
  23. Mayr, E.: 1954, 'Change of Genetic Environment and Evolution', in J. S. Huxley, A. C. Hardy and E. B. Ford (eds.), Evolution as a Process, Allen & Unwin, London, pp. 157-180.Google Scholar
  24. Mayr, E.: 1963, Animal Species and Evolution, Harvard University Press, Cambridge, MA.Google Scholar
  25. Nijhout, H. F.: 1991, The Development and Evolution of Butterfly Wing Patterns, Smithsonian Institution Press, Washington, DC.Google Scholar
  26. Nijhout, H. F.: 1994, 'Symmetry Systems and Compartments in Lepidopteran Wings: The Evolution of a Patterning Mechanism', Development Supplement, 225-233.Google Scholar
  27. Quine, W. V.: 1969, 'Natural Kinds', in W. V. Quine, Ontological Relativity and Other Essays, Columbia University Press, New York, pp. 114-138.Google Scholar
  28. Sober, E. and Lewontin, R. C.: 1982, 'Artifact, Cause and Genic Selection', Philosophy of Science 49, 157-180.Google Scholar
  29. Sober, E. and Wilson, D. S.: 1994, 'A Critical Review of Philosophical Work on the Units of Selection Problem', Philosophy of Science 61, 534-555.Google Scholar
  30. Szathmáry, E. and Maynard Smith, J.: 1997, 'From Replicators to Reproducers: The First Major Transitions Leading to Life', Journal of Theoretical Biology 187, 555-571.Google Scholar
  31. Wagner, G. P.: 1995, 'The Biological Role of Homologues: A Building Block Hypothesis', N. Jb. Geol. Paläont. Abh. 19, 279-288.Google Scholar
  32. Wagner, G. P.: 1996, 'Homologues, Natural Kinds and the Evolution of Modularity', American Zoologist 36, 36-43.Google Scholar
  33. Wagner, G. P. and Altenberg, L.: 1996, 'Perspective: Complex Adaptations and the Evolution of Evolvability', Evolution 50, 967-976.Google Scholar
  34. Williams, G. C.: 1966, Adaptation and Natural Selection, Princeton University Press, Princeton.Google Scholar
  35. Wimsatt, W. C.: 1980, 'Reductionistic Research Strategies and their Biases in the Units of Selection Controversy', in T. Nickles (ed.), Scientific Discovery, Volume II, Historical and Scientific Case Studies, Reidel, Dordrecht, pp. 213-259.Google Scholar
  36. Wimsatt, W. C.: 1981, 'The Units of Selection and the Structure of the Multi-level Genome', in P. D. Asquith and R. N. Giere (eds.), PSA 1980, Vol. 2, Philosophy of Science Association, East Lansing, MI, pp. 122-183.Google Scholar

Copyright information

© Kluwer Academic Publishers 1999

Authors and Affiliations

  • Robert N. Brandon
    • 1
  1. 1.Department of Philosophy and ZoologyDuke UniversityDurhamU.S.A. E-mail

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