Bachelor, Lawrence University, Appleton, Wisconsin, 1962. Doctor of Science (honorary), Lawrence University, Appleton, Wisconsin, 1981. Doctor of Philosophy, Harvard University, 1966.
Postdoctoral fellow Harvard University, 1966—1967. Jane Coffin Childs postdoctoral fellow Medical Research Council Laboratory Molecular Biology, Cambridge, England, 1967—1970. Faculty Yale University, New Haven, since 1970, Sterling professor molecular biophysics and biochemistry.
Investigator Howard Hughes Medical Institute, Chevy Chase, Maryland, since 1986. Visiting professor University Colorado, Boulder, 1992—1993.
Fellow: American Academy Arts & Sciences (Newcomb Cleveland prize 2001). Member: National Academy of Sciences.