The former director of the Institute of Molecular and Cell Biology at the University of Porto. He spent much of his career working in the biology department of the University of California, Berkeley.
He studied Theoretical Physics in South Africa and received his Ph.D. in 1972 from the University of Paris.
He and his partner, journalist Richard Zimler, were married in 2010 when same-sex marriage was legalized in Portugal.
His father was also a prominent scientist and one of the first to study fungi.
He worked at the University of California, Berkeley for almost 18 years, the same school from which philosopher Richard Montague graduated.