in Coventry and then won a state scholarship to Corpus Christi College, Cambridge, where he gained a BA degree (first-class honours) in medical sciences in 1965, and promoted to an MA in 1969. He obtained his PhD degree in Physiological Optics at the University of California, Berkeley, in the United States, as a Harkness Fellow in 1968. From 1968 to 1979 he was a Demonstrator and then Lecturer in Physiology at the University of Cambridge, and was also Director of Medical Studies at Downing College. From 1976 to 1979 he held the Royal Society Locke Research Fellowship.