D. S. Falconer and Introduction to Quantitative Genetics

Abstract

Anecdotal, Historical and Critical Commentaries on Genetics Edited by James F. Crow and William F. Dove DOUGLAS Falconer died on February 23, 2004, in Edinburgh. In his career he made 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), which for many throughout the world has been both their lead into and their lifelong reference on the subject. The first edition was published in 1960 when one of us (W.G.H.) was an undergraduate student in agriculture and becoming interested in animal breeding; the book (albeit advanced for that naïve student) gave a lead into the science behind the practice. The fourth edition in 1996 was co-authored by T.F.C. Mackay (Falconer and Mackay 1996). We both have read it many times, consulted it, and taught from it. The book has been part of our lives. In this Perspectives we shall give some brief biographical information about Douglas, who was our teacher, colleague, and friend, and a summary of some of his most significant research. We shall concentrate, however, on the development and impact of Introduction to Quantitative Genetics. 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 Falconer (unpublished notes) wrote that Thompson presided over the final year course, “but did almost nothing. I asked him at the beginning for recommendations as to what to read and he said ‘Just browse, my boy, just browse.’ So I worked away on my own … and at the end of the year he came along to me and said ‘Well, Douglas, my boy, you’re a very good lad and I don't think we need give you an examination this year.'” But the Dean prevailed and Douglas had to take an exam, in which he received First Class Honors. 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; but Douglas did not find it inspiring. While still a student he met and married Margaret Duke, a classicist, and they had two 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