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Fellows Directory

Alan Grafen

Professor Alan Grafen FRS


Elected: 2011



ORCID 0000-0002-1843-6457


Alan Grafen is an evolutionary biologist who applies mathematical and logical theory to problems in evolution. He has improved our understanding of how signals function in nature, successfully producing a mathematical model of Zahavi’s handicap principle, which states that characteristics that at first may appear to impede — such as the peacock’s tail — can actually signal genetic superiority to a potential mate and therefore be of benefit.

Alan applies mathematics and logic to areas in evolution that include animal signals, sexual selection, kin selection and kin recognition. He leads the Formal Darwinism Project at the University of Oxford, aiming to capture Darwin’s central argument concerning the role of natural selection in evolution within a mathematical framework.

The ultimate goal of Alan’s current work is to develop an overarching theory justifying fitness optimisation — or maximising genetic potential — as a central biological concept. As well as many academic papers, Alan has published two well-received books: the textbook Modern Statistics for the Life Sciences (2002) and a 2006 book honouring the work of fellow evolutionary biologist Richard Dawkins.

Interests and expertise

Subject groups


Fitness maximisation, Fundamental theorem of natural selection, Natural selection, Price Equation