RUPERT BILLINGHAM - "Bill" to his friends and to his colleagues in the fields of immunology and organ transplantation - was born in Wiltshire, where his father was a fish merchant. Although he moved to the United States in 1957 together with his wife, Jean, and his first two children, he never lost his endearingly broad country accent, a fact that both puzzled and intrigued Americans. Perhaps he meant this to convey, with the ebullience that was typical of him: "Here I am and you'll jolly well just have to take me as you find me!" Well, they did, and he became a widely respected and admired member of the American immunological community.
In the Second World War Billingham saw active service as a Lieutenant on Royal Navy anti-submarine escort vessels in Western Approaches Command and on the East Indies Station, and he retained a lifelong interest in books about ships and the sea. After the war he returned to Oxford, to Oriel College, to complete his DPhil under the supervision of Peter Medawar, who in 1943 had embarked on investigations into the causes of the almost invariable destruction of skin grafts when these are transplanted from one individual to another of the same species.
Such grafts later became known as allografts. In one way or another Billingham devoted the whole of his scientific life to the study of allografts - the immunological causes of their rejection, how to prevent them from being rejected, and why it is that the mammalian foetus, which is to be regarded as a natural allograft because half the genes that determine the cellular molecules that trigger rejection in a foreign environment come from the father, is not normally rejected.
During the course of these studies he made many very outstanding contributions to the field of organ transplantation, which has made such headlong strides since he began to work in it. Billingham was undoubtedly one of its great pioneers.
Billingham moved to Birmingham when Medawar became Professor of Zoology there in 1947 and, together with Medawar and an endocrinologist, P.L. Krohn, he engaged in a groundbreaking study in rabbits that demonstrated for the first time that the life of skin allografts can be very significantly prolonged by steroid hormones - an observation that eventually found its way into the clinic in the treatment of transplant recipients.
However, in 1951 Medawar accepted the prestigious Jodrell Chair of Zoology at University College London and Billingham, now married, once again pulled up his roots and accompanied Medawar as a Senior Research Fellow. By this time he had established a unique experimental partnership with Medawar, who I suspect would not have made the move without him.
They took with them a young man who had just completed his Zoology degree, initially at Billingham's suggestion, and so it was that the writer of this obituary joined the research team that became known in the mid-1950s in American transplantation circles as "the holy trinity". (This was by dint of a stream of innovative scientific papers under the imprint of "Billingham, Brent and Medawar", always strictly in alphabetical order.) I think that this designation was meant to be complimentary. I had always imagined that Medawar was thought of as the Father and myself as the Son, but the mind boggles at the irreverent and iconoclastic Bill being cast in the role of the Holy Ghost . . .
The department in which Billingham found himself at UCL was both highly intellectual and left-wing, and he settled into it with his usual cheerful and no-nonsense manner. Medawar has described this decade as the most creative period of his professional life and Billingham was right at the centre of it. He was a superb experimentalist who matched Medawar's inspirational role, though this is not at all to imply that Billingham could not be highly innovative. He had a deeply questioning mind that was often applied to the solving of difficult technical problems. …