WTH the death of John Robert Stanley Fincham in February 2005, genetics lost one of its most thoughtful and talented practitioners. His many research contributions, together with his other writings, were a guiding framework for the field of fungal genetics and biology in the twentieth century.
John was born in 1926, attended Hertford Grammar School, and went on to Cambridge University where he graduated with a degree in botany in 1946. He stayed on to obtain his Ph.D. in 1950 for work on mutants of Neurospora crassa deficient in the assimilation of ammonium, working under David G. Catcheside. His last year of doctoral study was spent at the California Institute of Technology, where he met and married Ann Emerson, the daughter of pioneer fungal geneticist Sterling Emerson and the granddaughter of R. A. Emerson, one of the founders of maize genetics (NELSON 1993). The Finchams had one son and three daughters.
John was appointed first as lecturer in botany (19501954) and then as reader (1954-1960) at Leicester University. He moved on to become head of the genetics department of the John Innés Institute until 1966, taking leave during his first year to go to the Massachusetts Institute of Technology as visiting associate professor of genetics (FINCHAM 1988).
John was appointed as professor and head of the new department of genetics at Leeds University in 1966. After a decade at Leeds and a sabbatical year at Caltech, John moved to new pastures, first as Buchanan Professor of Genetics at the University of Edinburgh from 1976 to 1984 and then to Cambridge as Arthur Balfour Professor of Genetics from 1984 to 1991. He then "retired" and moved back to Edinburgh, but continued working actively and publishing until 2000. He was elected a fellow of the Royal Society in 1969 and was a member of its council from 1974 to 1976. In 1977, he received the Emil Christian Hansen medal (Copenhagen), and he was elected to the Royal Society of Edinburgh in 1978. John served as editor of Heredity from 1971 to 1978 and as president of The Genetical Society from 1978 to 1981.
John's greatest contributions were to fungal genetics. His work on the biochemical and molecular genetics of Neurospora was early, varied, and profound. He started this work in the late 1940s, shortly after Beadle and Tatum had developed the one gene, one enzyme hypothesis and the technical groundwork for modern research on the organism (see HOROWITZ 1991 and PERKINS 1992). John's thesis on animation-deficient mutants of Neurospora initiated a clear line of work that continued for the rest of his career.
His work on the am gene of Neurospora, which encodes the hexameric NADP-glutamate dehydrogenase, was fundamental to progress in several areas of biochemical genetics (see DAVIS 2000). The most important was the mechanism by which allelic mutations could complement one another. At the time, the debate over the nature of "pseudoalleles" (alleles that failed to complement, but nevertheless could recombine with one another) had been resolved by the powerful demonstrations of intragenic recombination in bacteriophage (BENZER 1955). However, the resulting new definition of the gene-a segment of DNA whose mutant forms could not complement one another-had become an article of faith. John's laboratory was among the first to demonstrate "intragenic complementation," a phenomenon that created quite a stir in the late 1950s. His discovery came in a roundabout way, through finding "pseudowild" progeny from am1 X am2 crosses (FiNCHAM and PATEMAN 1957). These progeny were complementing heterokaryons arising from disomic ascospores that in turn arose from meiotic nondisjunction of homologous chromosomes. John observed more directly that a number of allelic pairs of am mutants could complement one another in heterokaryons, despite the fact that only one gene and one enzyme were involved (FiNCHAM 1959). His group probed more deeply into the phenomenon, showing that complementation could take place in vitro. …