Professor Geoffrey Beale was one of the UK's leading protozoan geneticists with an international reputation. His enthusiasm for the use of genetic analysis to address fundamental questions in the biology of both free-living and parasitic protozoa was based on the influence of a series of leading geneticists whom he worked with in the early part of his career in the John Innes Horticultural Institute and Cold Spring Harbor. His research started with plants and Escherichia coli. Later he moved his research to Paramecium, under the mentorship of Tracy Sonneborn (ForMemRS 1964). His work on the surface antigens of Paramecium was published in the early 1950s, at a time when there was much debate about the role of nuclear genes and elements in the cytoplasm in determining phenotype. This was a period when the nature of the genetic material was uncertain and many scientists believed that genes were proteins. Two of his papers published at that time provided a synthesis that explained how environmental signals and the cytoplasmic state determined whether a gene was expressed (1, 2)*. These findings provided one of the first cases of the regulation of gene expression at a conceptual level and provided a bridge between those promoting cytoplasmic inheritance and those fixed on nuclear genes as the sole determinants of phenotype. In the period from 1966, he also initiated research on extranuclear (mitochondrial) genetics and on the genetics of rodent and human malaria parasites, leading to several fundamental and important findings. This was a very productive period and included the establishment of links with a malaria research group in Thailand directed by Dr Sodsri Thaithong, leading to studies on the genetic basis of malaria drug resistance and studies on strain variation in Plasmodium falciparum. In the latter part of his career and into retirement, he took a more advisory role in his research group but his enthusiasm and interest stimulated those around him enormously.
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