Alan Grafen

Professor of Theoretical Biology

Tutorial Fellow of St John’s College


Research Interests

The formal Darwinism project is an ambitious attempt to provide a mathematical population genetic justification for the concept of fitness optimisation, at a very high level of abstraction. As well as advancing the highly technical core of the project, I am also engaged in applications, including (i) inclusive fitness on networks, in relation to altruism and population viscosity (ii) reinterpreting bet-hedging in terms of fitness optimisation (iii) showing that group selection has no similar justification leading to a conclusion of optimisation of group fitness. This work also has a historical interest. It is essentially capturing Darwin’s central claim in the Origin of Species that the mechanical processes of inheritance and reproduction (now represented by population genetics) give rise to the appearance of design (now represented by optimisation). By incorporating all the advances in evolutionary theory (ESS theory, inclusive fitness, optimality theory), the end result will be a grand overarching theory justifying fitness optimisation as a central concept in biology. It is a mathematical version of the synthesis of theory and its grounding in fundamental concepts to be found in Richard Dawkins’ “The Selfish Gene”. I also engage from time to time in other topics in evolutionary theory and in statistics. In February 2014 I completed "phyreg", an R package, available from CRAN, that implements my 1989 Phylogenetic Regression.

Additional Information

I am the co-author with Rosie Hails of ‘Modern Statistics for the Life Sciences’ (2002, OUP), which introduces undergraduate students (and others!) to the powerful technique of General Linear Modelling. Online supplements show how to do all the exercises in Minitab, SPSS and SAS. More details on my website






Selected Publications

  • Nee, S, N. Colegrave, S.A. West and A. Grafen. 2005. ‘The illusion of invariant quantities in life histories’. Science 309: 1236-1239 (Profiled in: de Jong, Is Invariance Across Animal Species Just an Illusion?, Science 309: 1193-1195)
  • Grafen, A. and M. Ridley (editors), 2006. Richard Dawkins: how a scientist changed the way we think. Oxford University Press.
  • Grafen A, 2006. Optimization of inclusive fitness, Journal of theoretical Biology 238, 541-563
  • Grafen, A. 2007. Detecting kin selection at work using inclusive fitness. Proc. R. Soc. B 274: 713-719.
  • Grafen, A. 2007. The formal Darwinism project: a mid-term report. Journal of evolutionary Biology 20, 1243-1254.