Sydney Brenner, biologist

He shared the Nobel Prize in medicine in 2002 for his contribution to work unravelling how genes control cell division.

He and two colleagues, John Sulston and Robert Horvitz, traced a transparent roundworm known as C. elegans to determine how cells divide and create something new.

The findings on programmed cell death were key to understanding how cancers develop and laid the groundwork for making C. elegans a major model organism in research.

His most important contribution to science, however, was the work he did with Francis Crick, the co-discoverer of DNA, and others to determine the genetic code.

In 1961 they demonstrated that DNA is made up of a series of three nucleotides called codons, which encode the amino acids that make up a protein.

He also helped discover messenger RNA, the molecule that directs the cell’s production of amino acids.

Born in South Africa in 1927, Brenner spent much of his early career in Britain, earning his PhD from Oxford.

Later, he joined Cambridge University and shared an office with Crick for 20 years.

In the early 1990s, Brenner went to California where he first worked at the Scripps Research Institute in La Jolla and later rejoined Crick as a distinguished professor at the Salk Institute.

He spent the last part of his career building the biomedical sciences in Singapore, where he became its first honorary citizen.

Brenner is survived by his three children. His wife died in 2010.