My main interest is evolutionary biochemistry and bioenergetics, and specifically the process of chemiosmosis, through which cells generate energy in the form of ATP by way of proton gradients across membranes. Called ‘the most counter-intuitive idea in biology since Darwin’, the mechanism of chemiosmosis has been elucidated at atomic resolution, yet its evolutionary significance has received little attention.

My research focuses on three major transitions in evolution: the origin of life itself; the origin of the eukaryotic cell; and the evolution of fundamental traits shared by all eukaryotic cells, notably sex, sexes, speciation and senescence. I am exploring the hypothesis that energetics played a critical role in each transition, addressing specific questions theoretically, and by mathematical modelling, chemical and biochemical experiments.