1968, PhD in Biochemistry, Cambridge. 1951-56, with Dragon School, Oxford; 1956-60, with Magdalen College School, Oxford; since 1961, with Clare College, Cambridge. With Cancer Research UK: 1990-2010, head of a research laboratory; currently, emeritus member of the faculty. Fellow, Royal Society, London; Foreign Associate, US National Academy of Sciences. Member, Scientific Council, European Research Council. Nobel Laureate, Physiology or Medicine (2001).