His contributions to statistical human genetics included: the first methods using maximum likelihood for the estimation of human linkage maps; pioneering methods for estimating human mutation rates; the first estimates of mutation rate in humans (2 × 10−5 mutations per gene per generation for the X-linked haemophilia gene); and the first notion that there is a "cost of natural selection". He was the first to estimate the rate of human mutation in his 1932 book entitled The Causes of Evolution. At the John Innes Horticultural Institution, he developed the complicated linkage theory for polyploids; and extended the idea of gene-enzyme relationships with the biochemical and genetic study of plant pigments.
his contributions to statistical human genetics included the first methods using maximum likelihood for the estimation of human linkage maps; pioneering methods for estimating human mutation rates; the first estimates of mutation rate in humans (2 × 10−5 mutations per gene per generation for the x-linked haemophilia gene); and the first notion that there is a "cost of natural selection" he was the first to estimate the rate of human mutation in his 1932 book entitled the causes of evolution at the john innes horticultural institution he developed the complicated linkage theory for polyploids; and extended the idea of gene-enzyme relationships with the biochemical and genetic study of plant pigments