Whippets with a homozygous deletion were apparently less able runners although their overall appearance was significantly more muscular.
He has applied his methods to the identification and mapping of genes affecting economically important single-gene (e.g. polled, double-muscling, callipyge, weaver, congenital muscular distonia), as well as complex multi-gene traits (e.g. milk and fattening yield and quality, fertility, disease resistance).