| Citation | Distinguished for his theoretical and experimental studies of population genetics and evolutionary biology. He extended the theory of selection in age-structured populations which provides the genetical foundation for the study of the evolution of life-history patterns and ageing, and demonstrated experimentally genetic variation in such traits. He made major contributions to the theory of the related topics of selection for the rate of genetic recombination, the evolution of complexes of tightly linked genes and the evolution of separate sexes and outbreeding, and showed experimentally that recombination rates could be changed by selection. He has developed a theory for the population dynamics of transposable genetic elements which provide standard models for the analysis and interpretation of data on transposable element frequencies. |