Advertisement

Evolution is About Populations, But Its Causes are About Individuals

  • Pierrick BourratEmail author
Original Article

Abstract

There is a tension between, on the one hand, the view that natural selection refers to individual-level causes, and on the other hand, the view that it refers to a population-level cause. In this article, I make the case for the individual-level cause view. I respond to recent claims made by McLoone that the individual-level cause view is inconsistent. I show that if one were to follow his arguments, any causal claim in any context would have to be regarded as vindicating a form of population-level cause view. I show why this is implausible and how a consistent individual-level cause position can be held within the interventionist account of causation. Finally, I argue that there is one sense in which natural selection might be said to refer to population-level causes of evolutionary change. The upshot is that, as noted by others, natural selection can be regarded as referring to a population-level cause in the context of frequency-dependent selection and other situations of fitness-altering interactions between the individuals of a population. But whether this statement is true will depend on the empirical case investigated, not some a priori conceptual distinction. Thus, even though situations of frequency dependence might be ubiquitous, it is orthogonal to the conceptual question of whether frequency-independent natural selection—McLoone’s target—refers to individual- or population-level causes.

Keywords

Causation Drift Individual-level causes Modularity Natural selection Population-level causes 

Notes

Acknowledgements

I am thankful to Mathieu Charbonneau, Paul Griffiths, Jun Otsuka, Peter Takacs, and two anonymous reviewers for comments on previous versions of the manuscript. I also thank the members of the Theory and Method in Biosciences group at the University of Sydney and in particular Stefan Gawronski who proofread the final manuscript. This research was supported by a Macquarie University Research Fellowship and a Large Grant from the John Templeton Foundation (Grant ID 60811).

References

  1. Abrams M (2009) Fitness “kinematics”: biological function, altruism, and organism–environment development. Biol Philos 24(4):487–504.  https://doi.org/10.1007/s10539-009-9153-2 CrossRefGoogle Scholar
  2. Ayala FJ, Campbell CA (1974) Frequency-dependent selection. Annu Rev Ecol Syst 5:115–138CrossRefGoogle Scholar
  3. Birch J (2017) The philosophy of social evolution. Oxford University Press, OxfordCrossRefGoogle Scholar
  4. Bouchard F, Rosenberg A (2004) Fitness, probability and the principles of natural selection. Br J Philos Sci 55(4):693–712CrossRefGoogle Scholar
  5. Bourrat P (2015a) Distinguishing natural selection from other evolutionary processes in the evolution of altruism. Biol Theory 10(4):311–321CrossRefGoogle Scholar
  6. Bourrat P (2015b) How to read ‘heritability’ in the recipe approach to natural selection. Br J Philos Sci 66(4):883–903CrossRefGoogle Scholar
  7. Bourrat P (2016) Generalizing contextual analysis. Acta Biotheor 64(2):197–217CrossRefGoogle Scholar
  8. Bourrat P (2017) Explaining drift from a deterministic setting. Biol Theory 12(1):27–38CrossRefGoogle Scholar
  9. Bourrat P (2018) Natural selection and drift as individual-level causes of evolution. Acta Biotheor.  https://doi.org/10.1007/s10441-018-9331-1 CrossRefGoogle Scholar
  10. Bourrat P (2019a) In what sense can there be evolution by natural selection without perfect inheritance? Int Stud Philos Sci 32(1):39–77CrossRefGoogle Scholar
  11. Bourrat P (2019b) Natural selection and the reference grain problem. Stud Hist Philos Sci A.  https://doi.org/10.1016/j.shpsa.2019.03.003 CrossRefGoogle Scholar
  12. Bourrat P (forthcoming) Causation and SNP heritability. Philos SciGoogle Scholar
  13. Coulson T, Benton T, Lundberg P, Dall S, Kendall B, Gaillard JM (2006) Estimating individual contributions to population growth: evolutionary fitness in ecological time. Proc R Soc B 273(1586):547–555.  https://doi.org/10.1098/rspb.2005.3357 CrossRefGoogle Scholar
  14. Dieckmann U, Ferrière R (2004) Adaptive dynamics and evolving biodiversity. In: Ferrière R, Dieckmann U, Couvet D (eds) Evolutionary conservation biology. Cambridge University Press, Cambridge, pp 188–224.  https://doi.org/10.1017/CBO9780511542022.015 CrossRefGoogle Scholar
  15. Fisher RA (1930) The genetical theory of natural selection: a complete variorum edition. Oxford University Press, OxfordCrossRefGoogle Scholar
  16. Frank SA (1998) Foundations of social evolution. Princeton University Press, PrincetonGoogle Scholar
  17. Frank SA (2014) The inductive theory of natural selection: summary and synthesis. arXiv:14121285 [physics, q-bio] 1412.1285
  18. Glennan S (2009) Productivity, relevance and natural selection. Biol Philos 24(3):325–339.  https://doi.org/10.1007/s10539-008-9137-7 CrossRefGoogle Scholar
  19. Godfrey-Smith P (2009) Darwinian populations and natural selection. Oxford University Press, OxfordCrossRefGoogle Scholar
  20. Griffiths PE, Pocheville A, Calcott B, Stotz K, Kim H, Knight R (2015) Measuring causal specificity. Philos Sci 82(4):529–555.  https://doi.org/10.1086/682914 CrossRefGoogle Scholar
  21. Hitchcock C, Velasco J (2014) Evolutionary and Newtonian forces. Ergo 1(2):39–77Google Scholar
  22. Huneman P (2012) Natural selection: a case for the counterfactual approach. Erkenntnis 76:171–194CrossRefGoogle Scholar
  23. Illari P, Russo F (2014) Causality: philosophical theory meets scientific practice. Oxford University Press, OxfordGoogle Scholar
  24. Krimbas CB (2004) On fitness. Biol Philos 19:185–203CrossRefGoogle Scholar
  25. Lande R, Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210–1226CrossRefGoogle Scholar
  26. Lewontin RC (1974) Annotation: the analysis of variance and the analysis of causes. Am J Hum Genet 26:400Google Scholar
  27. Luque VJ (2017) One equation to rule them all: a philosophical analysis of the Price equation. Biol Philo 32(1):97–125.  https://doi.org/10.1007/s10539-016-9538-y CrossRefGoogle Scholar
  28. Lynch KE, Bourrat P (2017) Interpreting heritability causally. Philos Sci 84(1):14–34CrossRefGoogle Scholar
  29. Matthen M, Ariew A (2002) Two ways of thinking about fitness and natural selection. J Philos 99:55–83CrossRefGoogle Scholar
  30. Matthen M, Ariew A (2009) Selection and causation. Philos Sci 76:201–224CrossRefGoogle Scholar
  31. McLoone B (2018) Why a convincing argument for causalism cannot entirely eschew population-level properties: discussion of Otsuka. Biol Philos 33(1–2):11.  https://doi.org/10.1007/s10539-018-9620-8 CrossRefGoogle Scholar
  32. Millstein RL (2006) Natural selection as a population-level causal process. Br J Philos Sci 57(4):627–653.  https://doi.org/10.1093/bjps/axl025 CrossRefGoogle Scholar
  33. Millstein RL (2013) Natural selection and causal productivity. In: Chao HK, Chen ST, Millstein RL (eds) Mechanism and causality in biology and economics. Springer, Dordrecht, pp 147–163CrossRefGoogle Scholar
  34. Nowak MA, Tarnita CE, Wilson EO (2010) The evolution of eusociality. Nature 466:1057–1062CrossRefGoogle Scholar
  35. Okasha S (2006) Evolution and the levels of selection. Oxford University Press, OxfordCrossRefGoogle Scholar
  36. Okasha S (2016) The relation between kin and multilevel selection: an approach using causal graphs. Br J Philos Sci 67(2):435–470CrossRefGoogle Scholar
  37. Otsuka J (2016a) Causal foundations of evolutionary genetics. Br J Philos Sci 67(1):247–269.  https://doi.org/10.1093/bjps/axu039 CrossRefGoogle Scholar
  38. Otsuka J (2016b) A critical review of the statisticalist debate. Biol Philos 31(4):459–482.  https://doi.org/10.1007/s10539-016-9528-0 CrossRefGoogle Scholar
  39. Pearl J (2009) Causality: models, reasoning, and inference, 2nd edn. Cambridge University Press, New YorkCrossRefGoogle Scholar
  40. Pence CH, Ramsey G (2013) A new foundation for the propensity interpretation of fitness. Br J Philos Sci 64(4):851–881.  https://doi.org/10.1093/bjps/axs037 CrossRefGoogle Scholar
  41. Pocheville A, Griffiths PE, Stotz K (2017) Comparing causes – an information-theoretic approach to specificity, proportionality and stability. In: Leitgeb H, Niiniluoto I, Sober E, Seppälä P (eds) Proceedings of the 15th Congress of Logic, Methodology and Philosophy of Science, College Publications, London, pp 260–275Google Scholar
  42. Price GR (1970) Selection and covariance. Nature 227:520–21CrossRefGoogle Scholar
  43. Ramsey G (2006) Block fitness. Stud Hist Philos Sci C 37:484–498Google Scholar
  44. Ramsey G (2016) The causal structure of evolutionary theory. Australas J Philos 94(3):421–434.  https://doi.org/10.1080/00048402.2015.1111398 CrossRefGoogle Scholar
  45. Reisman K, Forber P (2005) Manipulation and the causes of evolution. Philos Sci 72(5):1113–1123.  https://doi.org/10.1086/508120 CrossRefGoogle Scholar
  46. Rice SH (2004) Evolutionary theory: mathematical and conceptual foundations. Sinauer, SunderlandGoogle Scholar
  47. Rosenberg A, Bouchard F (2005) Matthen and Ariew’s obituary for fitness: reports of its death have been greatly exaggerated. Biol Philos 20:343–353CrossRefGoogle Scholar
  48. Russo F (2009) Causality and causal modelling in the social sciences: measuring variations. Methodos Series. Springer, DordrechtCrossRefGoogle Scholar
  49. Sarkar S (2008) A note on frequency dependence and the levels/units of selection. Biol Philos 23(2):217–228.  https://doi.org/10.1007/s10539-007-9092-8 CrossRefGoogle Scholar
  50. Sesardic N (2005) Making sense of heritability. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  51. Shapiro L, Sober E (2007) Epiphenomenalism—the do’s and the don’ts. In: Wolters G, Machamer P (eds) Thinking about causes: from Greek philosophy to modern physics. University of Pittsburgh Press, Pittsburgh, pp 235–264Google Scholar
  52. Snijders TAB, Bosker RJ (1999) Multilevel analysis: an introduction to basic and advanced multilevel modeling. SAGE, LondonGoogle Scholar
  53. Sober E (2013) Trait fitness is not a propensity, but fitness variation is. Stud Hist Philos Sci C 44(3):336–341.  https://doi.org/10.1016/j.shpsc.2013.03.002 CrossRefGoogle Scholar
  54. Sober E, Wilson DS (1998) Unto others: the evolution and psychology of unselfish behavior. Harvard University Press, CambridgeGoogle Scholar
  55. Spirtes P, Glymour CN, Scheines R (2000) Causation, prediction, and search, vol 81. MIT press, CambridgeGoogle Scholar
  56. Steel D (2006) Methodological individualism, explanation, and invariance. Philos Soc Sci 36(4):440–463.  https://doi.org/10.1177/0048393106293455 CrossRefGoogle Scholar
  57. Steel D (2008) Across the boundaries: extrapolation in biology and social science. Environmental ethics and science policy series. Oxford University Press, OxfordGoogle Scholar
  58. Stephens C (2004) Selection, drift, and the “forces” of evolution. Philos Sci 71(4):550–570.  https://doi.org/10.1086/423751 CrossRefGoogle Scholar
  59. Walsh DM (2000) Chasing shadows: natural selection and adaptation. Stud Hist Philos Sci C 31(1):135–153Google Scholar
  60. Walsh DM (2010) Not a sure thing: fitness, probability, and causation. Philos Sci 77(2):147–171CrossRefGoogle Scholar
  61. Walsh DM, Lewens T, Ariew A (2002) The trials of life: natural selection and random drift. Philos Sci 69(3):429–446CrossRefGoogle Scholar
  62. Walsh DM, Ariew A, Matthen M (2017) Four Pillars of Statisticalism. Philosophy and Theory in Biology 9(20170609),  https://doi.org/10.3998/ptb.6959004.0009.001
  63. Waters CK (2007) Causes that make a difference. J Philos 104(11):551–579CrossRefGoogle Scholar
  64. Woodward J (2003) Making things happen: a theory of causal explanation. Oxford University Press, New YorkGoogle Scholar
  65. Woodward J (2010) Causation in biology: stability, specificity, and the choice of levels of explanation. Biol Philos 25(3):287–318CrossRefGoogle Scholar
  66. Woodward J (2016) Causation and manipulability. In: Zalta EN (ed) The Stanford encyclopedia of philosophy, winter 2016 edn. https://plato.stanford.edu/archives/win2016/entries/causation-mani/

Copyright information

© Konrad Lorenz Institute for Evolution and Cognition Research 2019

Authors and Affiliations

  1. 1.Department of PhilosophyMacquarie UniversityNorth RydeAustralia
  2. 2.Department of Philosophy & Charles Perkins CentreUniversity of SydneySydneyAustralia

Personalised recommendations