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Mutual policing and repression of competition in the evolution of cooperative groups


EVOLUTIONARY theory has not explained how competition among lower level units is suppressed in the formation of higher-level evolutionary units1,2. For example, the key problem of early evolution is how small, individual replicators formed cooperative groups of sufficient complexity to allow accurate copying of the genetic material3. The puzzle is why parasites did not subvert the formation of cells by obtaining benetics benefits from the group without contributing to shared traits that enhance reproduction4. These parasites would outcompete other replicators within the cell, disrupting reproductive fairness among subunits and destroying the functional coherence of the group. A similar problem arose at a later evolutionary stage with the orderly mendelian segregation of subunits (chromosomes) within cells, and reproductive fairness continued to be a problem in the evolution of insect5 and human societies6. Here I present a simple model to show how reproductive fairness evolves among subunits to create functional coherence and higher-level units. Self-restraint, which evolves according to the kin-selection coefficient of relatedness, is not sufficient: mutual policing and enforcement of reproductive fairness are also required for the evolution of increasing social complexity.

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  1. Wilson, D. S. & Sober, E. J. theor. Biol. 136, 337–356 (1989).

    Article  CAS  Google Scholar 

  2. Maynard Smith, J. & Szathmáry, E. The Major Transitions in Evolution (Freeman, New York, 1995).

    Google Scholar 

  3. Eigen, M. & Schuster, P. The Hypercycle: A Principle of Natural Self-Organization (Springer, New York, 1979).

    Book  Google Scholar 

  4. Maynard Smith, J. Nature 280, 445–446 (1979).

    Article  Google Scholar 

  5. Ratnieks, F. L. W. & Reeve, H. K. J. theor. Biol. 158, 33–65 (1992).

    Article  Google Scholar 

  6. Alexander, R. D. The Biology of Moral Systems (Aldine de Gruyter, New York, 1987).

    Google Scholar 

  7. Frank, S. A. Proc. R. Soc. Lond. B 258, 153–161 (1994).

    Article  ADS  CAS  Google Scholar 

  8. Bremermann, H. J. & Pickering, J. J. theor. Biol. 100, 411–426 (1983).

    Article  CAS  Google Scholar 

  9. Lloyd, W. F. Two Lectures on the Checks to Population (1833, reprinted by Augustus M.Kelley, New York, 1968).

    Google Scholar 

  10. Hardin, G. Science 162, 1243–1248 (1968).

    Article  ADS  CAS  Google Scholar 

  11. Hardin, G. Living within Limits: Ecology, Economics and Population Taboos (Oxford Univ. Press, 1993).

    Google Scholar 

  12. Ratnieks, F. L. W. & Visscher, P. K. Nature 342, 796–797 (1989).

    Article  ADS  Google Scholar 

  13. Hurst, L. D. Proc. R. Soc. Lond. B 258, 287–298 (1994).

    Article  ADS  Google Scholar 

  14. Wilson, D. S. & Sober, E. Behav. Brain Sci. 17, 585–684 (1994).

    Article  Google Scholar 

  15. Bonner, J, T. The Cellular Slime Moulds 2nd edn (Princeton Univ. Press, 1967).

    Google Scholar 

  16. Morrissey, J. H. in The Development of Dictyostelium discoideum (ed. Loomis, W. F.) 411–449 (Academic, New York, 1982).

    Book  Google Scholar 

  17. Williams, J. G. et al. Cell 59, 1157–1163 (1989).

    Article  CAS  Google Scholar 

  18. Ozaki, T. et al. Development 117, 1299–1308 (1993).

    CAS  PubMed  Google Scholar 

  19. Maynard Smith, J. Evolution and the Theory of Games (Cambridge University Press, Cambridge, 1982).

    Book  Google Scholar 

  20. Price, G. R. Nature 227, 520–521 (1970).

    Article  ADS  CAS  Google Scholar 

  21. Hamilton, W. D. Nature 228, 218–220 (1970).

    Article  Google Scholar 

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Frank, S. Mutual policing and repression of competition in the evolution of cooperative groups. Nature 377, 520–522 (1995).

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