AB, Stanford University, 1947. Master of Arts, Stanford University, 1948. Doctor of Philosophy, Stanford University, 1951.
Biologist, National Institutes of Health, Bethesda, Maryland., 1953-1961; associate biochemist, Massachusetts General Hospital, Boston, 1961-1969; assistant professor microbiology, Harvard Medical School, Boston, 1966-1975; associate professor, Harvard Medical School, Boston, 1975-1982; research director, Boston biomedical Research Institute, 1967-1982; research professor division biological science, U. Montana, Missoula, since 1982. Director Stella Duncan Research Institute, Missoula, since 1982. Consultant Miles Laboratory, Elkhart, Indiana, 1980-1984.
Member American Association for the Advancement of Science (president Pacific division 1984-1985), American Society for Microbiology (National Foundation for Microbiology lecturer 1970, divisional lecturer 1978), American Society Biological Chemists.