A student of Dobzhansky, his work on genetic loads and on average effects of new mutants when in heterozygote was widely noticed. He has had a long career at Cornell University and at Virginia Polytechnic University, where he became University Distinguished Professor Emeritus. He died in 2015.
He is author of a number of books, including Radiation, Genes and Man (with Theodosius Dobzhansky), Genetic Load, Its Biological And Conceptual Aspects, The Search For The Gene, and The Study of Gene Action (with Joseph Falkinham).
Obituary at the Virigina Tech webpages
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