In Memoriam: Professor Richard Carter FRSE (1945-2021)

13 Oct 2021

Professor Richard Carter, an RSTMH Life Fellow and winner of our Chalmers Medal, who also published in our journal, Transactions, passed away after a short illness on 4 September 2021.

Richard was born on 26 February 1945 in Sydney, Australia, the second of three sons of Harold and Mary Carter. His father Harold (BVSc, DVSc (Hon), FRSE, AM) was a veterinary research scientist and historian. His mother Mary was a psychiatrist. In his youth the family lived on their small sheep farm at Ulundri, 20 miles outside the city.

When he was nine years old the family moved to Scotland where Richard entered school first in the village of Penicuik, then at George Watson’s College in Edinburgh. In 1967 he obtained his BSc in Biochemistry at the University of Edinburgh. Richard’s research began when he joined the Protozoan Genetics Research Group, in the Institute of Animal Genetics, also at Edinburgh University. Here he studied under the supervision of Geoffrey Beale FRS.

Richard’s initial work aimed to reveal the genetic organisation of Paramecium, but he was soon attracted to a highly imaginative project “hatched” by Geoffrey and P.C.C. Garnham FRS. to study the genetic organisation of malaria parasites. On this he worked with Dr David Walliker a parasitologist who had been recruited from Garnham’s laboratory. Together they built the foundations for our current understanding of malarial genetics. Using rodent malaria parasites, they showed Plasmodium followed Mendelian principles; is haploid in its blood stages, and that genetic recombination occurred (following sexual fertilisation) when it passes through the mosquito. Coincidentally their need to work with clones revealed many available parasite isolates were mixtures, an observation that provided a new clarity to the taxonomy of Plasmodium spp.

The malaria life cycle

From 1974 to 1986 he studied at the National Institutes of Health (NIH), Washington under the leadership of Dr “Lou” Miller. This proved to be the most productive period of his scientific life, he enjoyed intellectual freedom, excellent collaborators, and generous resources. The published foundations of malaria genetics were eventually validated for human malaria (Plasmodium falciparum) and applied to study questions such as drug resistance.

But Richard, with a background in genetics, inevitably became fascinated by the amazing biology of the sexual stages of the malarial life cycle. Richard’s deep interest in the history of his chosen research led him to investigate intriguing data published by Don Eyles and Clay Huff from the 1950s, suggesting immunisation with mixed blood stage parasites blocked transmission to the mosquito. Richard clearly demonstrated this inhibition resulted from an immune response to the sexual stages and was mediated by antibodies, complement and (in later collaborations with K. Mendis’ lab) cytokines.

He showed that immunisation with purified gametes can reduce transmission by >99%. Both Richard and his colleague Bob Gwadz, showed this immunity could be long-lived and rapidly boosted by blood infection. This phenomenon, now called transmission blocking immunity (TBI). is the foundation of an international effort to develop innovative new malaria vaccines (TBV).

In subsequent collaborative biochemical and field studies, he identified key target antigens of TBI, of which protein P230 was prominent. He then demonstrated this protein alone could induce antibodies that blocked fertilisation.

Chalmers Medal

His team also began an important body of work over many years on diverse aspects of the molecular regulation of sexual development both in the human host and in the mosquito. In 1985 for his collected body of work he was awarded the Chalmers medal by RSTMH.

In 1986 he returned to his old department in Edinburgh, initially as a member of the MRC external staff, then from 2008, as Professor of Parasite Genetics. Here his team developed high-throughput genetic methodologies, notably the technique of Linkage Group Selection (LGS) to identify the genetic mechanisms by which the parasite naturally evades either antimalarial drugs or host strain-specific protective immunity.

But his long-held fascination with Pfs230 continued. He modelled the structure/functional interactions of the "230 protein family" and how its members might act to bind the gametes together. His hand-drawn pictures (he was an excellent draftsman) remain a masterpiece of creative thinking and today continue to provide the foundation for ongoing efforts to develop 230 as a leading TBV candidate.

Intellectual maverick

He further developed his epidemiological interests in the human malarias. In collaborative studies in PNG and notably Sri Lanka he modelled the spatial dynamics of transmission and the effective implementation of transmission reduction measures.

In his later career he expanded his interest in the evolution of human malarias, notably of the “vivax type” (of which he had previously obtained unfortunate first-hand data on P. cynomolgi infection). In 2017 this contribution was recognised by the naming of a new member of the species complex P.carteri.

In 2007 he was elected as a Fellow of the Royal Society of Edinburgh.

Richard was an intellectual maverick, his wide-ranging body of work was invariably; conceptually and methodologically original, and the historical foundations of his work deeply-researched. He always questioned dogma and was prepared to think outside the box. He was a fount of knowledge and an inspiration not only to his many students and post docs, but to the wider community. Unsurprisingly his hypotheses sometimes generated controversy, but it must be said they were usually correct.

A generous colleague

As a colleague I can say he was generous in acknowledging the contributions of others, but above all he was justifiably secure in the value of his own work.

As an historian of malaria research, his memoirs on the development of the ideas underpinning malaria genetics and transmission blocking immunity provide an illuminating and enjoyable read.

Richard died after a short illness on 4 September 2021 and is survived by his wife Margaret with whom he shared a love of music and art – Richard was a talented pianist and painter.

He was instinctively creative, whether it was building a scientific hypothesis, a fish pond in the garden or toys for his children and grandchildren. He shared with his three sons, an enchanting world of stories called “The Perfumed Space Garden”, a land they fondly recall to this day, occupied by heroes, princes, princesses, ogres and Baron Butzenshoosoff.

He was also enthusiastic in sharing his interest and deep concern for the natural world with his grandchildren, keeping it simple and always fun-loving. His creativity will be missed by us all.