Biology & Philosophy

, Volume 29, Issue 2, pp 225–232 | Cite as

Adaptation, fitness and the selection-optimality links

  • Samir OkashaEmail author
  • Cédric Paternotte


We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we compare Grafen’s project with Fisher’s ‘fundamental theorem of natural selection’, and we speculate about whether Grafen’s results can be extended to a game-theoretic setting.


Formal Darwinism Grafen Optimality Fitness Selection Adaptation 



Thanks to Alan Grafen, Warren Ewens and Andy Gardner for discussion. SO was supported by the European Research Council Seventh Framework Program (FP7/2007–2013), ERC Grant agreement no. 295449. CP was supported by the Alexander von Humboldt Foundation.


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Copyright information

© Springer Science+Business Media Dordrecht 2014

Authors and Affiliations

  1. 1.University of BristolBristolUK
  2. 2.LMU UniversityMunichGermany

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