| Citation | Distinguished for his novel studies on the organisation and rearrangement of immunoglobulin genes (particularly human), which provided major new insights into the way the immune system works. He collaborated with others to estimate V-gene diversity by molecular hybridisation methods. He cloned globin cDNA, demonstrated splicing of nuclear RNA into kappa-chain mRNA, discovered that gene deletions underlie the Ig class switch and, with others, the repetitive sequences involved. He showed that in humans the kappa chains are encoded by fewer than 50 germ-line genes and established the arrangement of the heavy chain genes. He mapped the kappa and lambda genes and provided evidence for the involvement of the Ig locus in chromosomal translocations in Burkitt's lymphoma and the Philadelphia chromosome. |