Skip to main content

The formal darwinism project in outline

Abstract

The formal darwinism project aims to provide a mathematical framework within which important fundamental ideas in large parts of biology can be articulated, including Darwin's central argument in The Origin (that mechanical processes of inheritance and reproduction can give rise to the appearance of design), modern extensions of evolutionary theory including ESS theory and inclusive fitness, and Dawkins' synthesis of them into a single structure. A new kind of argument is required to link equations of motion on the one hand to optimisation programs on the other, and a major point is that the biologist's concept of fitness maximisation is not represented by concepts from dynamical systems such as Lyapunov functions and gradient functions. The progress of the project so far is reviewed, though with only a brief glance at the rather complicated mathematics itself, and the centrality of fitness maximisation ideas to many areas of biology is emphasised.

This is a preview of subscription content, access via your institution.

References

  • Barrett R, Rogers SM, Schluter D (2008) Natural selection on a major armor gene in threespine stickleback. Science 322:255–257

    Article  Google Scholar 

  • Batty CJK, Crewe P, Grafen A, Gratwick R (in press). An individual fitness maximization principle in a class-structured population under uncertainty. J Math Biol

  • Cosmides LM, Tooby J (1981) Cytoplasmic inheritance and intragenomic conflict. J Theor Biol 89:83–129

    Article  Google Scholar 

  • Darwin CR (1859) The Origin of Species. John Murray, London

    Google Scholar 

  • Davies NB, Krebs JR, West SA (2012) An introduction to behavioural ecology. Wiley-Blackwell, London

    Google Scholar 

  • de Groot MH (1970) Optimal statistical decisions. McGraw Hill, New York

    Google Scholar 

  • Dempster E (1955) The maintenance of genetic heterogeneity. Quant Biol 20:25–32

    Article  Google Scholar 

  • Edwards AWF (1994) The fundamental theorem of natural selection. Biol Rev 69:443–474

    Article  Google Scholar 

  • Ellner S, Rees R (2006) Integral projection models for species with complex demography. Am Nat 167:410–428

    Article  Google Scholar 

  • Engen S, Lande R, Saether B-E (2009) Reproductive value and fluctuating selection in an age-structured population. Genetics 183:629–637

    Article  Google Scholar 

  • Engen S, Lande R, Saether B-E, Dobson FS (1981) Reproductive value and the stochastic demography of age-structured populations. Am Nat 174:795–804

    Article  Google Scholar 

  • Ewens WJ (1989) An interpretation and proof of the fundamental theorem of natural selection. Theor Popul Biol 36:167–180

    Article  Google Scholar 

  • Ewens WJ (2004) Mathematical population genetics I. Theoretical introduction. Springer, Berlin

    Book  Google Scholar 

  • Fisher RA (1918) The correlation between relatives on the supposition of mendelian inheritance. Trans R Soc Edinburgh 52:399–433

    Article  Google Scholar 

  • Fisher RA (1930) The genetical theory of natural selection. Oxford University Press. See Fisher (1999) for a version in print

  • Fisher RA (1999) The genetical theory of natural selection. Oxford University Press. A Variorum edition of the 1930 and 1958 editions, edited by JH Bennett

  • Fisher RA (1958) The genetical theory of natural selection. 2nd ed. Dover Press, Mineola

    Google Scholar 

  • Frank SA (1995) George Price’s contributions to evolutionary genetics. J Theor Biol 175:373–388

    Article  Google Scholar 

  • Frank SA (1998) The foundations of social evolution. Princeton University Press, Princeton

    Google Scholar 

  • Frank SA, Slatkin M (1990) Evolution in a variable environment. Am Nat 136:244–260

    Article  Google Scholar 

  • Gardner A, Grafen A (2009) Capturing the superorganism: a formal theory of group adaptation. J Evol Biol 22:659–671

    Article  Google Scholar 

  • Gardner A, West SA, Barton NH (2007) The relation between multilocus population genetics and social evolution theory. Am Nat 169:207–226

    Article  Google Scholar 

  • Grafen A (1984) Natural selection, kin selection and group selection. In: Krebs JR, Davies NB (eds) Behavioural ecology, 2nd ed. Blackwell Scientific Publications, Oxford, UK, pp. 62–84

    Google Scholar 

  • Grafen A (1999) Formal Darwinism, the individual-as-maximising-agent analogy, and bet-hedging. Proc R Soc Series B 266:799–803

    Article  Google Scholar 

  • Grafen A (2000) Developments of Price’s Equation and natural selection under uncertainty. Proc R Soc Ser B 267:1223–1227

    Article  Google Scholar 

  • Grafen A (2002) A first formal link between the Price equation and an optimisation program. J Theoret Biol 217:75–91

    Article  Google Scholar 

  • Grafen A (2003) Fisher the evolutionary biologist. J R Stat So Series D (The Statistican) 52:319–329

    Article  Google Scholar 

  • Grafen A (2006a) Optimisation of inclusive fitness. J Theor Biol 238:541–563

    Article  Google Scholar 

  • Grafen A (2006b) A theory of Fisher’s reproductive value. J Math Biol 53:15–60 doi:10.1007/s00285-006-0376-4

    Article  Google Scholar 

  • Grafen A (2007a) Detecting kin selection at work using inclusive fitness. Proc R Soc Series B 274:713–719

    Article  Google Scholar 

  • Grafen A (2007b). The formal Darwinism project: a mid-term report. J Evol Biol 20:1243–1254

    Article  Google Scholar 

  • Grafen A (2007c) Inclusive fitness on a cyclical network. J Evol Biol 20:2278–2283

    Article  Google Scholar 

  • Grafen A (2008) The simplest formal argument for fitness optimization. J Genet 87:421–433

    Article  Google Scholar 

  • Hamilton WD (1964) The genetical evolution of social behaviour. J Theor Biol 7:1–52

    Article  Google Scholar 

  • Houston AI, McNamara J (1999) Models of Adaptive Behaviour: An Approach Based on State. Cambridge University Press, Cambridge

    Google Scholar 

  • Kennedy JS (1992) The New Anthropomorphism. Cambridge University Press, Cambridge

    Book  Google Scholar 

  • Killingback T, Bieri J, Flatt T (2006) Evolution in group-structured populations can resolve the tragedy of the commons. Proc R Soc Series B 273:1477–1481

    Article  Google Scholar 

  • Kirkpatrick M, Johnson T, Barton N (2002) General models of multilocus evolution. Genetics 161:1727–1750

    Google Scholar 

  • Lande R (2007) Expected relative fitness and the adaptive topography of fluctuating selection. Evolution 61:1835–1846

    Article  Google Scholar 

  • Lande R (2008) Adaptive topography of fluctuating selection in a Mendelian population. J Evol Biol 21:1096–1105

    Article  Google Scholar 

  • Lloyd D (1977) Genetic and phenotypic models of natural selection. J Theor Biol 69:543–560

    Article  Google Scholar 

  • Manolio T, Collins FS, Cox NJ, Goldstein DB, Hindorff LA, Hunter DJ, McCarthy MI, Ramos EM, Cardon LR, Chakravarti A, Cho JH, Guttmacher AE, Kong A, Kruglyak L, Mardis E, Rotimi CN, Slatkin M, Valle D, Whittemore AS, Boehnke M, Clark AG, Eichler EE, Gibson G, Haines JL, Mackay TFC, McCarroll SA, Visscher PM (2009) Finding the missing heritability of complex diseases. Nature 461:747–753

    Article  Google Scholar 

  • Maynard Smith J (1978) Optimization theory in evolution. Ann Rev Ecol System 9:31–56

    Article  Google Scholar 

  • Maynard Smith J, Price GR (1973) The logic of animal conflict. Nature 246:15–18

    Article  Google Scholar 

  • McGregor PK, Krebs JR, Perrins CM (1981) Song repertoires and lifetime reproductive success in the great tit (Parus major). Am Nat 118:149–159

    Article  Google Scholar 

  • Meszéna G, Kisdi E, Dieckmann U, Geritz S, Metz J (2001) Evolutionary optimisation models and matrix games in the unified perspective of adaptive dynamics. Selection 2:193–210

    Article  Google Scholar 

  • Moran PAP (1964) On the nonexistence of adaptive topographies. Ann Hum Genet 27:383–393

    Article  Google Scholar 

  • Ohtsuki H, Hauert C, Lieberman E, Nowak M (2006) A simple rule for the evolution of cooperation on graphs and social networks. Nature 441:502–505

    Article  Google Scholar 

  • Postma E, Spyrou N, Rollins L, Brooks R (2011) Sex-dependent selection differentially shapes genetic variation on and off the guppy Y chromosome. Evolution 65:2145–2156

    Article  Google Scholar 

  • Price GR (1970) Selection and covariance. Nature 227:520–521

    Article  Google Scholar 

  • Price GR (1972) Extension of covariance selection mathematics. Ann Hum Genet 35:485–490

    Article  Google Scholar 

  • Ridley M (1983) The explanation of organic diversity: the comparative method and adaptations for mating. Clarendon Press, Oxford

    Google Scholar 

  • Rousset F (2004) Genetic structure and selection in subdivided populations. Princeton University Press, Princeton, New Jersey

    Google Scholar 

  • Segestråle U (2000) Defenders of the truth. Oxford University Press, New York

    Google Scholar 

  • Taylor PD (1990) Allele-frequency change in a class-structured population. Am Nat 135:95–106

    Article  Google Scholar 

  • Taylor PD (1996) Inclusive fitness arguments in genetic models of behaviour. J Math Biol 34:654–674

    Article  Google Scholar 

  • Taylor PD, Frank SA (1996) How to make a kin selection model. J Theoret Biol 180:27–37

    Article  Google Scholar 

  • Tinbergen N (1963) On aims and methods of ethology. Zeitschrift für Tierpsychologie 20, 410–433. This journal was renamed Ethology in 1986. The paper has been reprinted as Tinbergen (1985)

  • Tinbergen N (1985) On aims and methods of ethology. Animal Biology 55, 297–321. This journal was renamed Ethology in 1986. The paper is a facsimile of Tinbergen (1963)

  • van Doorn G (2009) Intralocus sexual conflict. Ann N Y Acad Sci 1168:52–71

    Article  Google Scholar 

  • West S, Griffin A, Gardner A, Diggle S (2006) Social evolution theory for microbes. Nat Rev Microbiol 4:597–607

    Article  Google Scholar 

  • West SA (2009) Sex allocation. Princeton University Press, Princeton

    Google Scholar 

  • Williams GC (1966) Adaptation and natural selection. Princeton University Press, Princeton

    Google Scholar 

  • Wright S (1969–1978) Evolution and the genetics of populations (Four Volumes). University of Chicago Press, Chicago

Download references

Acknowledgments

I am very grateful to the editors and to all those colleagues who will write commentaries on this article. I am also grateful to Richard Gratwick and Paul Crewe for robust and relevant discussions over the past year, and to the Research Centre of St John’s College, Oxford, for a grant (to AG and to Prof. Charles Batty) to pursue the project. Paul Crewe, Cécile Fabre, Andy Gardner, Richard Gratwick, Jarrod Hadfield, Tom Kemp, Samir Okasha, Stu West and Greg Wyatt commented very helpfully on an earlier draft.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Alan Grafen.

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Grafen, A. The formal darwinism project in outline. Biol Philos 29, 155–174 (2014). https://doi.org/10.1007/s10539-013-9414-y

Download citation

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10539-013-9414-y

Keywords

  • Formal darwinism
  • Fitness maximisation
  • Behavioural ecology
  • Biological design
  • Population genetics
  • Natural selection