Models in the Biological Sciences

  • Elisabeth A. Lloyd
Part of the Springer Handbooks book series (SHB)


Evolutionary theory may be understood as a set of overlapping model types, the most prominent of which is the natural selection model, introduced by Charles Darwin and Alfred Russel Wallace. Many of the most prominent models today are represented through mathematical population genetics, in which genetical representations of populations evolve over time to produce evolutionary change. I review the variety of evolutionary models – from genic to group to species selection models – and how they are confirmed through evidence today. I discuss both applications to cases where we do not know the genetics, and to animal behavior and evolution.


State Space Evolutionary Theory Natural Kind Evolutionary Stable Strategy Semantic View 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

evolutionary stable strategy


  1. 1.
    E.A. Lloyd: The nature of Darwin’s support for the theory of natural selection, Philos. Sci. 50, 112–129 (1983)CrossRefGoogle Scholar
  2. 2.
    J.R. Griesemer: Presentations and the status of theories. In: PSA 1984, Vol. 1, ed. by P.D. Asquith (Philosophy of Science Association, East Lansing 1984) pp. 102–114Google Scholar
  3. 3.
    B.C. Van Fraassen: The Scientific Image (Oxford Univ. Press, Oxford 1980)CrossRefGoogle Scholar
  4. 4.
    E.A. Lloyd: Units and levels of selection. In: Stanford Encyclopedia of Philosophy, ed. by E.N. Zalta (Stanford Univ., Stanford 2005),, Winter 2012 edn.Google Scholar
  5. 5.
    J. Beatty: What’s wrong with the received view of evolutionary theory? In: PSA 1980, Vol. 2, ed. by P.D. Asquith, R.N. Giere (Philosophy of Science Association, East Lansing 1980) pp. 397–426Google Scholar
  6. 6.
    P. Thompson: The structure of evolutionary theory: A semantic perspective, Stud. Hist. Philos. Sci. 14, 215–229 (1983)CrossRefGoogle Scholar
  7. 7.
    J. Beatty: Chance and natural selection, Philos. Sci. 51, 183–211 (1984)CrossRefGoogle Scholar
  8. 8.
    E.A. Lloyd: A semantic approach to the structure of population genetics, Philos. Sci. 51, 242–264 (1984)MathSciNetCrossRefGoogle Scholar
  9. 9.
    E.A. Lloyd: The Structure and Confirmation of Evolutionary Theory (Princeton Univ. Press, Princeton 1994)Google Scholar
  10. 10.
    P. Suppes: Models of data. In: Logic, Methodology, and Philosophy of Science, ed. by E. Nagel, P. Suppes, A. Tarski (Stanford Univ. Press, Stanford 1962) pp. 252–261Google Scholar
  11. 11.
    R. Giere: Explaining Science: A Cognitive Approach (Univ. Chicago Press, Chicago 1988)CrossRefGoogle Scholar
  12. 12.
    C. Alsina, R.B. Nelsen: Math made Visual: Creating Images for Understanding Mathematics (Mathematics Association of America, New York 2006)CrossRefzbMATHGoogle Scholar
  13. 13.
    H. Meyer-Ortmanns, S. Thurner (Eds.): Principles of Evolution (Springer, Berlin, Heidelberg 2011)Google Scholar
  14. 14.
    R. Dawkins: The Extended Phenotype (Oxford Univ. Press, Oxford 1982)Google Scholar
  15. 15.
    E.A. Lloyd, R.C. Lewontin, M. Feldman: The generational cycle of state spaces and adequate genetical representation, Philos. Sci. 75, 140–156 (2008)MathSciNetCrossRefGoogle Scholar
  16. 16.
    R.C. Lewontin: The Genetic Basis of Evolutionary Change (Columbia Univ. Press, New York 1974)Google Scholar
  17. 17.
    T. Dobzhansky: Genetics of the Evolutionary Process (Harper Rox, New York 1970)Google Scholar
  18. 18.
    R. Lewontin: Population genetics, Annu. Rev. Genet. 1(1), 37–70 (1967)CrossRefGoogle Scholar
  19. 19.
    R. Levins: Evolution in Changing Environments (Princeton Univ. Press, Princeton 1968)Google Scholar
  20. 20.
    M. Kimura, T. Ohta: Protein polymorphism as a phase of molecular evolution, Nature 229, 467–469 (1971)CrossRefGoogle Scholar
  21. 21.
    J.H. Gillespie: Population Genetics: A Concise Guide, 2nd edn. (Johns Hopkins Univ. Press, Baltimore 2004)Google Scholar
  22. 22.
    E. Mayr: Evolutionary challenges to the mathematical interpretation of evolution. In: Mathematical Challenges to the Neo-Darwinian Interpretation of Evolution, ed. by P.S. Moorehead, M.M. Kaplan (Wistar Institute, Philadelphia 1967) pp. 47–54Google Scholar
  23. 23.
    M. Wade: A critical review of the models of group selection, Q. Rev. Biol. 53, 101–114 (1978)CrossRefGoogle Scholar
  24. 24.
    M.J. Wade: Adaptation in Metapopulations (Univ. Chicago Press, Chicago 2016)CrossRefGoogle Scholar
  25. 25.
    E.A. Lloyd: Confirmation of evolutionary and ecological models, Biol. Philos. 2(3), 277–293 (1987)CrossRefGoogle Scholar
  26. 26.
    R.A. Skipper Jr.: The heuristic role of Sewall Wright’s 1932 adaptive landscape diagram, Philos. Sci. 71(5), 1176–1188 (2004)MathSciNetCrossRefGoogle Scholar
  27. 27.
    P. Forber: On biological possibility and confirmation, unpublished manuscript (2008)Google Scholar
  28. 28.
    P. Forber: Confirmation and explaining how possible, Stud. Hist. Philos. Sci. Part C: Stud. Hist. Philos. Biol. Biomed. Sci. 41(1), 32–40 (2010)CrossRefGoogle Scholar
  29. 29.
    L.A. Dugatkin, H.K. Reeve: Behavioral ecology and levels of selection: Dissolving the group selection controversy, Adv. Study Behav. 23, 101–133 (1994)CrossRefGoogle Scholar
  30. 30.
    K. Sterenly: Explanatory pluralism in evolutionary biology, Biol. Philos. 11(2), 193–214 (1996)CrossRefGoogle Scholar
  31. 31.
    E. Sober, D.S. Wilson: Unto Others: The Evolution and Psychology of Unselfish Behavior, (Harvard Univ. Press, Cambridge 1998), p. 57, pp. 98–99 Google Scholar
  32. 32.
    K. Sterenly, P.E. Griffiths: Sex and Death: An Introduction to Philosophy of Biology (Univ. Chicago Press, Chicago 1999) pp. 168–169, p. 172Google Scholar
  33. 33.
    B. Kerr, P. Godfrey-Smith: Individualist and multi-level perspectives on selection in structured populations, Biol. Philos. 17(4), 477–517 (2002)CrossRefGoogle Scholar
  34. 34.
    C.K. Waters: Why genic and multilevel selection theories are here to stay, Philos. Sci. 72, 311–333 (2005)CrossRefGoogle Scholar
  35. 35.
    R.D. Alexander, G. Borgia: Group selection, altruism, and the levels of organization of life, Annu. Rev. Ecol. Syst. 9, 449–474 (1978)CrossRefGoogle Scholar
  36. 36.
    M.K. Uyenoyama, M.W. Feldman: Evolution of altruism under group selection in large and small populations in fluctuating environments, Theor. Popul. Biol. 17, 380–414 (1980)CrossRefzbMATHGoogle Scholar
  37. 37.
    D.S. Wilson: The Natural Selection of Populations and Communities (Benjamin Cummings, Menlo Park 1980)Google Scholar
  38. 38.
    R.K. Colwell: Evolution of female-biased sex ratios: The essential role of group selection, Nature 290, 401–404 (1981)CrossRefGoogle Scholar
  39. 39.
    J.F. Crow, K. Aoki: Group selection for a polygenetic behavioral trait: A differential proliferation model, Proc. Natl. Acad. Sci. USA 79, 2628–2631 (1982)CrossRefzbMATHGoogle Scholar
  40. 40.
    R.E. Michod: The theory of kin selection, Annu. Rev. Ecol. Syst. 13, 23–55 (1982)CrossRefGoogle Scholar
  41. 41.
    M.J. Wade: Soft selection, hard selection, kin selection, and group selection, Am. Nat. 125, 61–73 (1985)CrossRefGoogle Scholar
  42. 42.
    M. Smith: Evolutionary progress and levels of selection. In: The Latest on the Best: Essays on Evolution and Optimality, ed. by J. Dupre (MIT Press, Cambridge 1987)Google Scholar
  43. 43.
    D.C. Queller: Quantitative genetics, inclusive fitness, and group selection, Am. Nat. 139, 540–558 (1992)CrossRefGoogle Scholar
  44. 44.
    P. Godfrey-Smith, R.C. Lewontin: The dimensions of selection, Philos. Sci. 60, 373–395 (1993)CrossRefGoogle Scholar
  45. 45.
    M.W. Feldman, L.L. Cavalli-Sforza: Cultural and biological evolutionary processes, selection for a trait under complex transmission, Theor. Popul. Biol. 9(2), 238–259 (1976)MathSciNetCrossRefzbMATHGoogle Scholar
  46. 46.
    M.W. Feldman, L.L. Cavalli-Sforza: Cultural Transmission and Evolution: A Quantitative Approach (No. 16) (Princeton Univ. Press, Princeton 1981)Google Scholar
  47. 47.
    E. Jablonka, M.J. Lamb: Evolution in four dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life, Life and Mind: Philosophical Issues in Biology and Psychology (MIT Press, Cambridge 2005)Google Scholar
  48. 48.
    F. Suppe: What’s wrong with the received view on the structure of scientific theories?, Philos. Sci. 39, 1–19 (1972)CrossRefGoogle Scholar
  49. 49.
    F. Suppe: The Structure of Scientific Theories, 2nd edn. (Univ. Illinois Press, Urbana 1977)Google Scholar
  50. 50.
    F. Suppe: Understanding scientific theories: An assessment of developments, Philos. Sci. 67, S102–S115 (2000)CrossRefGoogle Scholar
  51. 51.
    S.M. Downes: The Importance of Models in Theorizing: A Deflationary Semantic View, PSA, Vol. 1 (Univ. Chicago Press, Chicago 1992) pp. 142–153Google Scholar
  52. 52.
    P. Godfrey Smith: The strategy of model-based science, Biol. Philos. 21, 725–740 (2006)CrossRefGoogle Scholar
  53. 53.
    A. Love: Theory is as theory does: Scientific practice and theory structure in biology, Biol. Theory 7(4), 325–337 (2012)CrossRefGoogle Scholar
  54. 54.
    S. Scheiner: Toward a conceptual framework for biology, Q. Rev. Biol. 85, 293–318 (2010)CrossRefGoogle Scholar
  55. 55.
    J.M. Smith: The Evolution of Sex (Cambridge Univ. Press, Cambridge 1978), No. 574.1 S5Google Scholar
  56. 56.
    A. Love: Rethinking the structure of evolutionary theory for an extended synthesis. In: Evolution – The Extended Synthesis, ed. by M. Pigliucci, G.B. Müller (MIT Press, Cambridge 2010) pp. 403–441CrossRefGoogle Scholar
  57. 57.
    M. Pigliucci, G.B. Müller: Elements of an extended evolutionary synthesis. In: Evolution – The Extended Synthesis, ed. by M. Pigliucci, G.B. Müller (MIT Press, Cambridge 2010) pp. 3–17CrossRefGoogle Scholar
  58. 58.
    J. Griesemer: Presentations and the status of theories, PSA: Proc. Bienn. Meet. Philos. Sci. Assoc. (1984) pp. 102–114Google Scholar
  59. 59.
    I. Lakatos: The Methodology of Scientific Research Programmes. Philosophical Papers, Vol. 1 (Cambridge Univ. Press, Cambridge 2011)zbMATHGoogle Scholar
  60. 60.
    T. Piavani: An evolving research programme: The structure of evolutionary theory from a Lakatosian perspective. In: The Theory of Evolution and Its Impact, ed. by A. Fasolo (Springer, Milan 2012) pp. 211–228CrossRefGoogle Scholar
  61. 61.
    E.A. Lloyd: Model robustness as a confirmatory virtue: The case of climate science, Stud. Hist. Philos. Sci. 49, 58–68 (2015)CrossRefGoogle Scholar
  62. 62.
    A. Grafen: Natural selection, kin selection and group selection. In: Behavioural ecology: An Evolutionary Approach, Vol. 2, ed. by J.R. Krebs, N.B. Davies (Blackwell Scientific Publications, Oxford 1984) pp. 62–84Google Scholar
  63. 63.
    J.M. Townsend-Mehler, F.C. Dyer, K. Maida: Deciding when to explore and when to persist: A comparison of honeybees and bumblebees in their response to downshifts in reward, Behav. Ecol. Sociobiol. 65, 305–312 (2011)CrossRefGoogle Scholar
  64. 64.
    R. Lewontin: Interdeme selection controlling a polymorphism in the house mouse, Am. Nat. 96, 65–78 (1962)CrossRefGoogle Scholar
  65. 65.
    G.A. Parker, J.M. Smith: Optimality theory in evolutionary biology, Nature 348, 27–33 (1990)CrossRefGoogle Scholar
  66. 66.
    E.A. Lloyd, M.W. Feldman: Commentary: Evolutionary psychology: A view from evolutionary biology, Psychol. Inq. 13(2), 150–156 (2002)CrossRefGoogle Scholar
  67. 67.
    C. Allen: Models, mechanisms, and animal minds, South. J. Philos. 52, 75–97 (2014)CrossRefGoogle Scholar
  68. 68.
    C. Buckner: Two approaches to the distinction between cognition and ’mere association’, Int. J. Comp. Psychol. 24(4), 314–348 (2011)Google Scholar
  69. 69.
    T.T. Hills: Animal foraging and the evolution of goal-directed cognition, Cogni. Sci. 30, 3–41 (2006)CrossRefGoogle Scholar
  70. 70.
    T. Hills, P.M. Todd, R.L. Goldstone: Search in external and internal spaces: Evidence for generalized cognitive search processes, Psychol. Sci. 19, 676–682 (2008)CrossRefGoogle Scholar
  71. 71.
    P.M. Todd, T.T. Hills, T.W. Robbins: Building a foundation for cognitive search. In: Cognitive Search: Evolution, Algorithms, and the Brain, Vol. 9, ed. by P.M. Todd, T.T. Hills, T.W. Robbins (MIT Press, Cambridge 2012) pp. 1–7, Strüngmann Forum ReportsGoogle Scholar
  72. 72.
    R.S. Olson, A. Hintze, F.C. Dyer, D.B. Knoester, C. Adami: Predator confusion is sufficient to evolve swarming behavior, J. R. Soc. Interface 10, 20130305 (2013), CrossRefGoogle Scholar
  73. 73.
    C. Reynolds: Flocks, herds, and schools: A distributed behavioral model, Comput. Graph. 21(4), 25–34 (1987)CrossRefGoogle Scholar

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© Springer-Verlag Berlin Heidelberg 2017

Authors and Affiliations

  1. 1.Department of History and Philosophy of Science and MedicineIndiana UniversityBloomingtonUSA

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