Geoffrey Herbert Beale, MBE, FRS, FRSE 11 June 1913 - 16 October 2009 Geoffrey Beale was recognized internationally as a leading protozoan geneticist with an all-absorbing love of genetics, stimulated in the early part of his career by either working with or meeting many of the key figures who laid the foundations of modern genetics in the 1930s and 1940s. His work on the genetics of the surface antigens of Paramecium provided a conceptual breakthrough in our understanding of the role of the environment, the cytoplasm and the expression of genes, and he continued his interest in the role of cytoplasmic elements in heredity through studies on both the endosymbionts and mitochondria of Paramecium. He pioneered the genetic analysis of parasitic protozoa with his work on Plasmodium, and this stimulated many other scientists to take a genetic approach with these experimentally challenging organisms. Geoffrey was born in Wandsworth, London, on 11 June 1913, the son of Herbert Walter Beale and Elsie Beale (née Beaton). His family included an elder brother (Hugh) and two younger sisters (Margaret and Joan). When he was about five years old the family moved to Wallington, Surrey, where he spent the rest of his childhood as well as staying there during his university undergraduate and postgraduate studies. He attended Sutton County School, Surrey from 1923 until he obtained his higher school certificates in mathematics, physics and chemistry in 1931. His main interest at that time was music and he briefly considered the possibility that he might make music his career and he became an accomplished pianist and organist. However, he was also fascinated by the idea of a career in biological research. There were no biological sciences taught at the county school and no one in his family had ever been to university. Most of his knowledge about biological sciences was obtained by reading and as a result of this, he resolved to go to university and take up biological research as a career, despite much parental advice about the advantages of working in a bank or becoming a tax inspector. He accepted a place at Imperial College, London and, after some difficulties he was able to get a loan from Surrey County Council to support his attendance. He also got a job as organist at Cheam Presbyterian Church in 1931 to provide much-needed funds. This source of income, together with the loan from the council and money he received from other small jobs, enabled him to finance his studies. To reduce costs, he lived at home and cycled from Wallington to Imperial College in South Kensington, a round trip of 20 miles each day. Although he originally registered for zoology, he switched to botany and in 1935 received a BSc from Imperial College with first-class honours. In his third year he attended a summer course on plant genetics given by staff of the John Innes Horticultural Institute with presentations by leading figures in the field and this, plus his undergraduate studies, fired an interest in genetics that stayed with him throughout his career. After graduating, he applied to the John Innes Institute as a volunteer but was soon offered a job by its temporary head, J. B. S. Haldane. This was a great opportunity, because the Institute was one of only two places in the UK for research in genetics, and some of the leading geneticists of the time (Haldane, Darlington, Scott-Moncrieff & others) were members of staff. The job enabled him to receive a stipend as well as work on a PhD degree, initially under Haldane’s supervision but later was nominally supervised by Darlington. He received the PhD in 1938 for his thesis ‘The genetics of Verbena’. From 1935 to 1940 he worked on different projects that were being investigated in the institute and met and became friends with Dan Lewis, K. Mather, Mick Callan and A. C. Fabergé with the latter stimulating his interest in Russian culture and language. Although all the staff at the institute had been registered as being in ‘reserved occupations’, he was 1 nevertheless called up to the army in 1941 to undertake military training. Shortly after this he was drafted into the Intelligence Corps (Field Security) as a Corporal. He joined the Army Base Unit and was sent to Russia in a convoy arriving in Archangel in November 1941. British troops and equipment were being sent to Russia to aid in the war against Germany, and Beale was instructed to facilitate the unloading of ships, seeing to it that their contents were speeded on their way to the interior for the Russian and Allied forces and liaising with the Russian authorities to ensure this happened. In May 1942 he was transferred to Murmansk and was promoted to the rank of Captain. He then spent a short period in London, before being posted to Finland as part of the Allied Control Commission, which was overseeing the Armistice between the Russians and the Finns. He had an important role in the negotiations, particularly given his knowledge of Russian. He describes (in his unpublished autobiographical notes), in his own laconic way, that his main achievement during the war was to learn Russian. However, the facts would suggest that his contribution was significant because he rose to the rank of Captain and, in 1947, was awarded an MBE for his military service. He always underplayed this award, suggesting that he received it for just being there, and in a letter to Dan Lewis he commented, ‘Doubtless thousands of others have been awarded an MBE too, but still it’s better than nothing, like a PhD.’ On completion of his military service, his job at the John Innes was still technically available to him, but he decided to take a post at Cold Spring Harbor (USA) and joined Milislav Demerec’s laboratory in 1946 to work on the rate of mutation of Escherichia coli from phage sensitivity to resistance. At this time many leading geneticists visited during the summer or attended the regular symposia, and so he was able to meet Luria, Lederberg, Delbrück, Sonneborn and others at a time when these scientists were laying the groundwork for microbial genetics. He was also friendly with Barbara McClintock and it is possible that his discussions with her aroused his interest in cytoplasmic inheritance. In 1947, he obtained a Rockefeller Fellowship to work with Sonneborn on Paramecium at Bloomington (USA) but this required that he return to the UK at the end of his stay. His return was made possible by Waddington who offered him a lectureship at the Institute of Animal Genetics in Edinburgh. Geoffrey had a fruitful stay in Bloomington, Indiana, and published several significant papers on Paramecium. In 1948 he moved to Edinburgh and continued this research on Paramecium with the immobilization surface antigens with the aim of elucidating the relationship between the genes and environment. Using a genetic approach and the unique features of the system, he showed the essential role of the cytoplasm in determining the expression of these immobilisation antigens. This led to one of the first models of the regulation of gene expression by the environment. These findings need to be considered in the context of knowledge at that time. The nature of the gene was unclear and a number of phenotypes were considered to be determined by non-nuclear plasmagenes with unconventional modes of inheritance. The genetic analysis undertaken by Beale allowed him to develop hypotheses that began to explain the phenomenon of antigenic variation as the product of the interaction between the gene, the cytoplasm and the environment and to move away from the concept of plasmagenes. In 1954 he published a monograph entitled ‘The genetics of Paramecium aurelia’ reviewing the current and past research on this organism together with a general introduction promoting the value of genetic analysis and the utility of Paramecium as an experimental system In 1963, based on his research, he was appointed as a Royal Society Research Professor and he held this position until his retirement in 1978. Based on the prestige of this appointment he was able to obtain funding from Edinburgh University and the Wellcome Trust, he was able to design and build The Protozoan Genetics Unit and developed a research group devoted to protozoan genetics. He continued his work on the immobilisation antigens in the subsequent 15 years with the characterization of the antigens and biochemical analysis of their synthesis, essentially provided one of the first examples of the regulation of gene expression by environmental conditions. The establishment of the Protozoan Genetics group allowed him to also expand his research into other areas, initially with Paramecium but subsequently with parasitic protozoa. Research in Sonneborn’s laboratory in the 1940s and 1950s had identified a series of Paramecium stocks that produced a substance or particle capable of killing other sensitive stocks. He started research on one class of particles, the mate killers that contained the symbiont (mu), so called because it was only during mating that they killed sensitive strains. The results from a series of experiments led to the metagon hypothesis which postulated that there were genetic elements in 2 the cytoplasm that determined the killer phenotype. These findings generated considerable scientific interest because they provided support for the concept that phenotype could be determined by non-nuclear genetic elements. Subsequently, a careful reinvestigation of the experiments showed that the basic observations on which the metagon theory was based were not repeatable, and the metagon hypothesis was reluctantly declared defunct by Beale himself. The work on symbionts as extranuclear genetic elements led Beale to the study of the genetic determinants of mitochondria.