Douglas Falconer

Douglas Falconer (March 101913 - February 23 2004), is mostely known for his many important contributions to quantitative genetics and to the genetics of the mouse. Undoubtedly, however, his greatest impact was through his textbook, Introduction to Quantitative Genetics (Falconer 1960a). The first edition was published in 1960, gave a lead into the science behind the practice in animal breeding. The fourth edition in 1996 was co-authored by T.F.C. Mackay (FALCONER and MACKAY 1996).

Biographical information

Douglas Falconer's family, which came from Edinburgh, had no significant tradition of science. His father was a minister in the church when Douglas was born on March 10, 1913, in Old Meldrum, Aberdeenshire, but the family soon moved back to Edinburgh, where Douglas went to school. There he developed an interest in science, although no biology was taught. He was accepted for university but his studies were delayed by 5 years when he contracted tuberculosis. During those long years of recuperation he read widely, including Morgan's Theory of the Gene. Finally, in 1936 he resumed his education at St. Andrews University, where he shone in science, being both more able and older than his contemporaries and, judging by his later years, a clearer thinker and more organized. He took an honors degree in zoology under D'Arcy Thompson (of Growth and Form fame), but did not study any genetics. Falconer (unpublished notes) wrote that Thompson presided over the final year course, "but did almost nothing. Douglas received first class honors at his exam. He was unfit for military service and took his Ph.D. in 1943 under James Gray at Cambridge on "The behavior of wireworms in relation to temperature and light", which is less esoteric than it appears as the wireworm is a major crop pest and there was a wartime need to increase agricultural production. Douglas did not find it inspiring.

While still a student he met and married Margaret Duke, a classicist, and they had three sons; all three survive him. Among their shared interests was a love of music, which they played together. Douglas played the flute and continued to do so until he was well into his eighties. In midlife Douglas developed diabetes, which he managed well, but he became increasingly blind in his last years. To those who met him from at least the 1960s onward, he already appeared a frail individual, but his frailness belied his determination and toughness. He was still coming into work and writing long after his retirement in 1980, not the least on further editions of his book, and he still kept an interest and enjoyed scientific discussion until his death.

Subsequent to completing his Ph.D., Douglas had a temporary lectureship at Queen Mary College London, then based in Cambridge, where his teaching included a course in genetics. At the end of the war, the Agricultural Research Council (ARC) was planning an institute for the study of genetics in relation to animal breeding and was bringing a group together, initially near London, but later to move to Edinburgh. Seeing this prospect, in 1945 Douglas took the opportunity to work with R. A. (Sir Ronald) Fisher at Cambridge (whose mouse labs were in his own house). Douglas set to work on mapping mutant genes and also to work on the inheritance of milk yield in mice (recording the trait as the increase in litter weight after feeding, not with a mechanical milking machine) but his time with Fisher was not entirely satisfactory. He recalls in an anecdote:

Fisher was engaged in a complicated experiment involving a large number of mouse genes, and also involving inbreeding, that is, brother by sister mating. Inbreeding is well known to reduce fertility, and I thought it would be interesting to see how much the fertility would be reduced in the next two or three generations, and see whether he would have sufficient animals to choose suitable pairs to mate, and I found out that he would not, that he would find that many of his lines would become extinct because there was not a suitable pair to continue them. I thought I ought to tell him of this finding and see what he thought about it, so I did one day at tea. He listened to what I said and then without uttering a word, he turned round and walked out of the room. The next day he came to me and said, "I think you should arrange to go to Edinburgh as soon as you can."

Douglas Falconer was appointed to the Genetics section of the ARC Animal Breeding and Genetics Research Organization in Edinburgh in 1947, under the direction of Professor C. H. Waddington, newly appointed to the Buchanan Chair of Animal Genetics at the University of Edinburgh. A very illustrious group were brought together. In addition to Falconer, those in quantitative genetics included Alan Robertson, Jim Rendel (who left in 1951 for Australia), Eric Reeve, Forbes Robertson, and Ian Mason and, later, Crad Roberts and George Clayton.Waddington essentially allowed people to work with little direction; in fact, the set up was much more like a modern university department with independently funded, self-directed groups, rather than the traditional hierarchical European university department.

Douglas Falconer wrote Introduction to Quantitative Genetics, the first edition of which was published in 1960 (FALCONER 1960a). In the preface he states his objectives:

My aim in writing this book has been to provide an introductory textbook of quantitative genetics, with the emphasis on general principles rather than on practical application, and one moreover that can be understood by biologists of no more than ordinary mathematical ability. In pursuit of this latter aim I have set out the mathematics in the form that I, being little of a mathematician, find most comprehensible, hoping that the consequent lack of rigour and elegance will be compensated for by a wider accessibility. The reader is not, however, asked to accept conclusions without proof... I have no particular class of reader in mind, but have tried to make the book useful to as wide a range of readers as possible (p. v).

The first edition of his book was a major success. It was reprinted five times by Oliver & Boyd and twice by Longman. Douglas realized the increasing need for a new edition, but other commitments prevented his giving time to it.

He had been appointed Deputy Director of the ARC Unit of Animal Genetics under Waddington, but "Wad" became less interested in genetics and also went on a long sabbatical. In 1968 Falconer was appointed to a Personal Chair in Genetics at the University of Edinburgh and also Director of the ARC Unit. The following year he was appointed Head of the Department of Genetics with responsibilities for all the diverse groups, including other major ones in mutation research, epigenetics, and protozoan genetics, housed in several buildings (later known as the Genetics Village) on the King's Buildings campus. This was a heavy task, which Douglas undertook in an even-handed manner, notwithstanding the competing claims of many prima donnas, giving it all the commitment he gave to other work including his research, which he continued throughout this time. In 1977 he was able to give up the Headship of the Department following John Fincham's appointment to the Buchanan Chair. He could then devote time to revising his book, both prior to his formal retirement and consequent closure of the ARC Unit of Animal Genetics in 1980 and subsequently when he continued research and writing but ceased active experimental work with the mouse.

Falconer's major contributions in quantitative genetics were on the response to artificial selection in mice, the concept of the cross-environment genetic correlation, and, as noted above, development of the theory for understanding the genetics of complex human diseases in terms of an underlying continuous liability. This research also provided much illustrative material for the book. Particularly in his early years of mouse genetic research, Falconer also worked on identification and mapping of individual genes and notably identified the first sex-linked gene in the mouse, Tabby (FALCONER 1952a).

Undoubtedly Douglas Falconer has had an enormous and lasting impact on quantitative genetics; indeed, he essentially defined the subject.