Bachelor of Science, Loyola U, Chicago, 1963; Master of Science, DePaul University, 1966; Doctor of Philosophy, Northwestern University, 1968.
National Institutes of Health fellow Michigan State University, East Lansing, 1968-1970. Assistant professor biology University of California at Los Angeles, 1969-1974, associate professor, 1974-1977, professor biology, 1977-1981, chairman biology, 1979-1981, dean, life science, 1981-1987. Vice chancellor for research, dean of graduate school, professor biology University North Carolina, professor biology Chapel Hill, 1988-1991.
Chancellor, professor biology University Pittsburgh, 1991-1995. Provost Smithsonian Institution, Washington, 1996-1999, under secretary for science, 2000—2002. Vice president research, dean graduate studies University Maryland, College Park, 2002—2004, professor biology, since 2004.
Visiting professor University Nijmegen, Netherlands, 1975-1976, Monash University, 1977.
Fellow American Association for the Advancement of Science. Member American Society Zoology, Society Development Biology, American Society Molecular Biology and Biochemistry, Business Higher Education Forum (vice chair 1994-1996, chair 1996-1998), Association Universities for Research in Astronomy (board directors).
Married, 1964; 3 children.