Education:

1961-1964

• B.S. in Chemistry, University of Pittsburgh, Pittsburgh, PA.
• Graduated Magna Cum Laude, Phi Beta Kappa.

1964-1969

• Ph.D. in Biochemistry, Department of Biochemistry, University of Wisconsin, Madison, WI.
• Supervisor: Dr. Julius Adler (Member of the National Academy of Sciences).
• Research: We were the first to identify the basal end of the bacterial flagellum, develop methods for its purification, and elucidate its fine structure and specific attachments to the cell envelopes of E. coli and B. subtilis.

Postdoctoral Training:

1970-1971

• Department of Biochemistry, University of Wisconsin, Madison, WI.
• Supervisor: Dr. William W. Cleland (Member of the National Academy of Sciences)
• Research: We synthesized the novel compound, Cr(III)ATP, and showed it to be a useful paramagnetic, dead-end inhibitor for exploring the active site and kinetic mechanisms of enzymes that required ATP as one of their substrates.

1971-1973

• Department of Biochemistry, Stanford University Medical School, Stanford, CA.
• Supervisor: Dr. Paul Berg (Nobel Laureate, Member of National Academy of Sciences)
• Research: Using the small DNA tumor virus, simian virus 40 (SV40), we were the first to develop a subcellular system that allowed animal virus DNA to continue replicating in vitro. We also applied somatic cell genetics to the study of SV40 replication in vivo.

Supervisory Positions:

• 1973-1978 Assistant Professor, Department of Biological Chemistry, Harvard Medical School, Boston, MA.
• 1978-1985 Associate Professor, Department of Biological Chemistry, Harvard Medical School, Boston, MA.
• 1985-1986 Professor (tenured), Department of Biological Chemistry, Harvard Medical School, Boston, MA.
• 1986-1995 Laboratory Head and Full Member (tenured) of the Roche Institute of Molecular Biology, Roche Research Center, Nutley, NJ.
• 1988-1995 Adjunct Professor, Department of Biological Sciences, Columbia University, New York, NY.
• 1996-present Section Chief (tenured) and member of the Senior Biomedical Research Service, National Institute of Child Health & Human Development, Bethesda, MD.

Publications:

225 primary research papers, reviews and books on chemistry, bacterial structure and physiology, enzymology, DNA replication, chromatin structure, and gene expression. These studies were carried out in vitro, and in bacteria, yeast, animal viruses, mammalian cells, frog eggs, mouse oocytes and preimplantation embryos.

Selected Awards and Patents: 

• 1964 Phi Beta Kappa Scholastic Honorary Fraternity
• 1974-1979 Established Investigator of the American Heart Association
• 1994-1995 Humboldt Senior Scientist Fellowship, University of Munich, Germany
• 2011 "NIH Director’s Challenge Award" to identify genes that regulate genome duplication in human cells
• 2022 “Autophagy modulators for use in treating cancer”, US patent #11,471,460
• 2022 "NICHD Lifetime Achievement Award"