Foundations of a mathematical theory of darwinism

Charles J.K. Batty, Paul Crewe, Alan Grafen, Richard Gratwick*

*Corresponding author for this work

Research output: Contribution to journalArticlepeer-review


This paper pursues the 'formal darwinism' project of Grafen, whose aim is to construct formal links between dynamics of gene frequencies and optimization programmes, in very abstract settings with general implications for biologically relevant situations. A major outcome is the definition, within wide assumptions, of the ubiquitous but problematic concept of 'fitness'. This paper is the first to present the project for mathematicians. Within the framework of overlapping generations in discrete time and no social interactions, the current model shows links between fitness maximization and gene frequency change in a class-structured population, with individual-level uncertainty but no uncertainty in the class projection operator, where individuals are permitted to observe and condition their behaviour on arbitrary parts of the uncertainty. The results hold with arbitrary numbers of loci and alleles, arbitrary dominance and epistasis, and make no assumptions about linkage, linkage disequilibrium or mating system. An explicit derivation is given of Fisher's Fundamental Theorem of Natural Selection in its full generality.

Original languageEnglish
Pages (from-to)295-334
Number of pages40
JournalJournal of Mathematical Biology
Issue number2
Early online date9 Jul 2013
Publication statusPublished - 31 Aug 2014


  • Fitness maximization
  • Formal darwinism
  • Price equation
  • Reproductive value


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