Copyright 0 1993 by the Genetics Society of America Perspectives

Anecdotal, Historical and Critical Commentaries onGenetics Edited by James F. Crow and William F. Dove

Quantitative Genetics in Edinburgh: 1947-1 980

Douglas Falconer

Institute of Cell, Animal and Population Biology, University ofEdinburgh, West Mains Road, Edinburgh, EH93JT, Scotland

OOD rationing in the second world war brought FISHER),and Edinburgh (with F. A. E. CREW).Edin- F home to everyonein Britain the need to improve burgh’s department was the first, established in 1919 agricultural output in order to reduce our depend- as the Animal Breeding Research Department, with ence on imports. Seeing clearly the need for more CREWas its Director but no otherstaff and no build- government-funded research on animal breeding, the ing. Aforceful and persuasive speaker,CREW ob- Agricultural Research Council (ARC) set up, in 1945, tained money from various sources toexpand the the Animal Breeding and Genetics Research Organi- department, and he cajoled several wealthy industri- zation (ABGRO). The “Genetics” in the title signified alists into providing funds for a new building and to the intention to pursuebasic genetics with experimen- endow achair. The Chair,founded in 1928 with tation on laboratory animals, to be done by the Ge- CREWas its first occupant, was called the Buchanan netics Section of ABGRO. R. G. WHITE,then Profes- Chair after Lord WOOLAVINGTONwhose family name sor of Agriculture in the University of North Wales was Buchanan and whose business was whisky distill- at Bangor, was appointed Director, with C. H. WAD- ing; half the funds needed for the endowment were DINGTON as Chief Geneticist in charge of the Genetics his gift. The title of the Chair was Animal Genetics, Section. WADDINGTON, then aged42, was preeminent but it was changed to Genetics in WADDINGTON’Stime. among the few geneticists in Britain at that time; his The new building was formally opened in 1930 with influential text, An Introduction to Modern Genetics, 12 scientific staff and 13 visiting researchers. had been published in 1939. Soon after his appoint- CREW’Senthusiasm attracted many visitors who mentto ABGRO, WADDINGTONwas offeredthe came for shortvisits or for longer periodsof research, Buchanan Chair of Animal Genetics in the University among whom were some notable figures-LANCELOT of Edinburgh in succession to F. A. E. CREW. This HOGBEN,JULIAN HUXLEY,J. B. S. HALDANE, and H. was the reason it was decided to locate ABGRO in J. MULLER(who was there from 1938 to 1940). On a Edinburgh. WADDINGTONthen held both positions, handsome oak panel in the entrance hall of the Insti- University Professor and Honorary Director of the tute building in gilded carved letters are thenames of Genetics Section of ABGRO. those who obtained higher degrees from the Depart- The Genetics Section, which had been in temporary ment. There are66 up to 1947, and 5 more by 1950 quarters in London, moved to Edinburgh in 1947. It when the inscriptions stopped. The first is CREW was housed together with the University Department himself who is recorded as obtaining a D.Sc. in 1921 in a building named the Institute of Animal Genetics. and a Ph.D. in 1923. Others who will be familiar to The main part of ABGRO was accommodated in a most geneticists are F. B. HUTT (Ph.D. 1929, D.Sc. large rented villa not far away until a new building on 1939), CHARLOTTEAUERBACH (Ph.D. 1935, D.Sc. the University campus close tothe Institute was 1947), and H. J. MULLER(D.Sc. 1940). opened in 1964. Under CREW’S leadershipthe Institute did pioneer- The location of ABGRO in Edinburgh continued a ing work on sex determination, reproductive physi- distinguished tradition of animal breeding and ge- ology, and many aspects of the husbandry and breed- netics there. There were then (I think) only three ing of sheep, cattle, pigs, horses, and poultry. There university departments of geneticsin the UnitedKing- was also work on cytology, on Drosophila genetics, dom:London’s University College (where J. B. s. and on the genetics of the color of budgerigars. (When HALDANEwas Professor),Cambridge (with R. A. I was a Ph.D. student in the Zoology Department in

Genetics 133: 137-142 (February, 1993) 138 D. Falconer

Cambridge I found a set of CREW’S budgerigarskins earlier work done by its members. Those in the Unit hidden away in a drawer. They made an impressive at the beginning in 1947 who worked on quantitative and beautiful illustration of all the main Mendelian genetics and related topics were the following: principles, and were amajor stimulus to my own C.H. WADDINGTON.His many diverse interests interest in genetics.) CREW’Sera culminated in the centered on developmental genetics. In quantitative holding of the Seventh InternationalCongress of Ge- genetics, he showed with Drosophila how what looked netics in Edinburgh in August, 1939. He was made superficially like Lamarckian inheritance of an ac- President in default of N. VAVILOVwho was unable quiredcharacter could result fromstraightforward to come (see the Perspectives of August, 1992). The selection. outbreak of the second world war brought the Con- ALANROBERTSON. With J. M. RENDEL he formu- gress and most of the Institute’sactivities to an abrupt lated improvement programs for dairy cattle,and the end. During the war CREW,who had amedical degree, “contemporarycomparison” by whichbulls are se- worked in the War Office on medical statistics. He lected for use in artificial insemination, which revo- resigned his Chair in 1944 because, so he said, he felt lutionized dairy cattle breeding. His experiments with himself to be too much out of date in genetics, but he Drosophila tested the adequacy of current selection returned to Edinburgh totake up the Chair of Public theory and located some of the genes responsible for Health and Social Medicine. the responses to selection, foreshadowing contempo- When the ARC group came to Edinburgh after the rary quantitative marker identification. Using KIMU- war, CREW’S Institute was much depleted in staff and RA’S stochastic theory, he developed a new theory of funds. Soon afterarrival in Edinburgh one of our selection limits in a finite population. technical staff, not renowned for his tact, found him- J. M RENDEL. Afterworking with ROBERTSONon self sitting next to an unknown person at coffee and, dairy cattle, he left in 1951 to join CSIROin Australia thinkingthat some conversation was called for, re- and became head of its genetics section. He iswell marked, “I understand that this place has been pretty known for his work on developmental canalization in inactive recently.” The unknown person was A. W. F. Drosophila. GREENWOODwho had been actingDirector during D. S. FALCONER.showed I that selection for growth CREW’Sabsence. It is no wonderthat those left of inmice was most effective when practiced in the CREW’Sstaff saw us newcomers as an arrogant lot environment in which the strain was expected to per- intent on an aggressive take-over. I fear that at first form (as opposed to the frequently advocatedpractice we were a sore trial to them. of selecting in the most favorable environment), and The ARC funded agricultural research in two main that a character measuredin two environments could ways. There were large groups in their own buildings be treated as two correlated characters. I introduced with a full-time director appointed by the ARC, and the use of realized heritability as a way of describing there were “units” which were small groups working selection response. in a university department under the direction of a R. A. BEATTY.In addition to studies of heteroploidy senior member of the university staff, usually the in mice and rabbits, he studied the genetics of sper- professor. ABGRO was a large group, but the Ge- matozoa, showing that metric characters of sperma- netics Section operated like a unit within it; its mem- tozoa are determined by the genotype of the testis bers were ABGRO staff but WADDINGTON,its direc- and not that of the spermatozoa. tor, was not. This anomalous situation was rectified F. W. ROBERTSON.In selecting for large and small in 1951 when H. P. DONALD,who had been in CREW’S body size in Drosophila he found strong asymmetry department, succeeded WHITE as Director of in the response to selection in the two directions, and ABGRO. The Genetics Section was then formally showed that at the selection limit there was still con- separated; ABGRO lost its G and became ABRO. In siderablegenetic variation. Chromosome assaysof 1957 the Genetics Section was designated the Unit of selected lines revealed strong epistatic interaction. Animal Genetics. Differences of body size were due to differences of In what follows I shall not discriminate between the cell number, not cellsize. Later he worked on the Genetics Section and the Unit, and will refer to both ecological and physiological genetics of Drosophila as the Unit. Itis about the Unit that I am writing here growth. In 1970he left fora chair in Aberdeen and I will not be able in this short article to say more University. about ABRO, though there was much fruitful collab- E. C. R. REEVE.He worked with F. W. ROBERTSON oration between the members of the two groups, and on selection in Drosophila and showed that inbred the work of ABRO was a large component of quanti- lines were considerably more variable than F1 hybrids. tative genetics in Edinburgh. Later he worked on bacterial genetics. To review adequately the work of the Unit would J. H. SANG. Hestudied population growth of Dro- be impossible. Instead I shall summarize briefly the sophila in culture, and developed a synthetic culture Perspectives 139 medium which became an essential tool for physiolog- it set forth were virtually unknown in the United ical genetics. He left for a Chair in the University of Kingdom despite the presence of HALDANEand Sussex in 1965. FISHERwho had provided much of the mathematical Some laterappointments in quantitative genetics background. There were few geneticists of any sort, were: and they tended to be regardedas eccentrics pursuing N. BATEMAN(1948). He selected for high and low anincomprehensible subject. Consequently little or milk production in mice and found very strong asym- no genetics was taught in undergraduate courses. metry of response. He transferred toABRO in 1957. Most of us, therefore, joined the Unit with very little I. L. MASON(1 949). He studied dual-purpose cattle background in genetics. Forexample, the nearest and advised on animal breeding programs in many thing to genetics in my zoology course at St. Andrews countries. He cataloged the origins and characteristics was the curious fact that Ascaris sheds mostof its of all livestock breeds. In 1972 he left to join FA0 in chromatin when it makes somatic cells. Some of us Rome. had not even a biological background; ALANROBERT- G. A. CLAYTON(1950). He worked with A. ROB- SON started as a physical chemist, and REEVEas a ERTSON on Drosophila selection and fitness experi- mathematician. After joiningthe unit, however, ROB- ments. They were,I think, the first to select with ERTSON spent nine months with SEWALLWRIGHT and replicate lines andto test theobserved responses J. L. LUSH,the two who had done most to develop against theoreticalpredictions. He also worked on quantitative genetics in its application to animal breed- turkeybreeding. He transferred to the University ing. Consequently,he was much betterinformed staff in 1959. about quantitative genetics and animal breeding than A.L. MCLAREN(1958). She studied maternal ef- the rest of us. In preparation for joining the Unit I fects, embryo transfer, early development, reproduc- spent 18 months with R. A. FISHERin Cambridge in tive physiology, and chimeras in mice. In 1974 she order to learn about mouse genetics. FISHERwas then left to be Director of the Medical Research Council’s mainly interested in linkage, and I did not learn much new Mammalian Development Unit in London. about quantitative genetics from him. R. C. ROBERTS(1959). He compared the life-time The original intention for the work on quantitative growth and reproduction of mouse lines selected for genetics was that there should be research on farm large and small body size and found that small mice animals (but without farm facilities), on rabbits, on had smaller litters, but more of them, thanlarge mice, mice, and on Drosophila. The basic quantitative ge- and produced nearly twice as many offspring in total. netics would be done with Drosophila because it is He characterized selected mouse lines using A. ROB- cheap and quick. But the results from Drosophila ERTSON’S theory of selection limits. could not be applieddirectly to farm animals because W. G. HILL,who was appointed to the University Drosophila is too different in physiology, in chromo- staff in 1965, must beincluded here because he some number, and in lacking crossing over in males. worked in close association with the Unit. His work The rabbits and mice were to form a bridge, being covered many aspects of theoretical quantitative ge- similar in physiology, chromosome number, and male netics, particularly in relation to selection andthe crossing over. Any breeding method that might be estimation of parameters. based on the Drosophila results would be tried with It was never the intention that thework of the Unit mice or rabbits and if it worked it could be applied shouldbe restricted to quantitative genetics. WAD- with more confidence to farm animals. It soon became DINGTON believed, as CREW had, that any aspect of apparent, however, that there was no great difference genetics might lead to advances in animal breeding. in the quantitative genetics of Drosophila and mice. Accordingly there were other members of the Unit So the chromosome number, male crossing over, and working on molecular genetics, cytology, develop- indeed the physiology, were largely irrelevant. ment, and systematics, among whom were H. G. CAL- No one“directed” our work. The ARC itself LAN who went to a chair at St. Andrews University in seemed to take no interest in what we did, or what we 1950 and J. L. SIRLIN fromabout 1962 to 1970. achieved. WADDINGTON,nominally our director, left We were generously provided with excellent tech- us free to do what we each thought best. This was a nical assistance. A great advantage of working in the wise policy, and it worked; I do not think that any of ARC. Unit was that funding was always assured; we us wasted much time in doing the wrong things. And did not have to spend time writinggrant applications. the freedom was greatly appreciated. By the 1940s, partly because of the war, Britain had The first experiments done with Drosophila and fallen farbehind theUnited States in quantitative mice were on selection. These take a long time, and genetics and the theory underlying animal breeding. when they finally produced results we were eager to In the UnitedStates, J. L.LUSH’S Animal Breeding publish them quickly. But in this we were frustrated. Plans had been published in 1937, but the principles We sent the papers to the Journal of Genetics, the 140 D. Falconer oldest of the two British journals publishing genetical affronted. I believe that the cool relationship between work. It was owned and edited by J. B. S. HALDANE, the Birmingham and Edinburgh schools that persisted who did the refereeing himself. But he was not well for many years may have had its origin in this unfor- organized. It was said that when he was away for some tunate episode. timethe cleaners, unwilling todisturb the pilesof The members of the ARC Unit, though ostensibly paper on his tables, covered them over with newspa- employed forresearch, contributed substantially to pers. When he returned he did not remove the papers, the Department’steaching. In respect of what we but started again on top. This was very useful to the actually did in the Institute, therewas little distinction geologists who, when HALDANEwas away, took their between Unit and University staff, and those not students to his room to demonstrate stratification. It familiar with the local arrangementsoften did not required several pleading letters of reminder before know to which group anyone belonged. The Univer- we eventually got our papers published. Continued sity authorities, however, were slow to recognize the difficulties with publication in British journals led existence of non-University staff and subjected us to WADDINGTONto found a new journal, Genetical Re- petty restrictions. For example, we could not be offi- search, in 1960.Edited by E. C. R. REEVE,it has cial supervisors of Ph.D. students; there had to be a flourished and earned a high reputation. University supervisor, usually WADDINGTONwho In 1948 I. M. LERNER,a visitor from California, often knew little or nothing of what the student was brought us new techniques from the United States. doing. Eventually, however, the authorities were per- He unfolded the mystery of SEWALL WRIGHT’S path suaded to be less narrow-minded. In 1949 a postgrad- coefficients, which were being used for deducing the uate Diploma in Genetics was started, allowing stu- necessary theoreticalparameters for quantitative dents with little previous training in genetics to em- breeding. LERNER wrote most of his Population Ge- bark on Ph.D. studies; it had a substantial component netics andAnimal Improvement, published in 1950, of quantitative genetics. Then, in 1975, an MSc. in while he was in Edinburgh. An important event in animal breeding was started in collaboration with the 1949 was a visit by SEWALL WRIGHTwho also spent a Agriculture Department and was run by W. G. HILL, sabbatical year in Edinburgh. I think, however, that who had worked on animal breeding and hadbecome his visit came too soon for some of us who did not a leading theorist in quantitative and population ge- have enough background to understand muchof what netics. This was the only course of its kind in the he had to teach us, though it was a useful stimulus in United Kingdom. It attracted students (aboutsix each showing us what a long way we had to go to catch up year) from all over the world, many of whom went on with current knowledge. He gave a long course of to take Ph.D.s in Edinburgh and then went home to lectures and these formed the basis for part of his found nuclei of quantitative genetics or animal breed- Evolution and the Genetics of Populations, the first vol- ing in their own countries, notably in Australia and ume of which was published in 1968. New Zealand. Not surprisingly, many of the staff of In 1950, near the end of WRIGHT’Svisit, a sympo- sium on quantitative inheritance was held in the Insti- ABRO or its successor organization were recruited tute; it was published in 1952. WRIGHTgave a lengthy from among our students. talk on the interactions between coat color genes in The Institute housed much more thanjust theARC guinea pigs. But the manuscript was lost on his way Unit. WADDINGTONrapidly increased the staff and home anda quite different paper appeared in the the range of research, and there were several groups published symposium. It was a synopsis of the current of workers with separate funding. The following is a state of quantitative genetics and was surely more very incomplete outline of the research staff of the generally useful than the guineapig paper would have Institute in the early 196Os, when the numbers were been. The symposium, however, had unforeseen and probably near their peak. The Unit then had nine regrettable consequences of a political nature. KEN- staff members, and the University 13, among whom NETH MATHER, then Professor of Genetics inBir- were the following. CHARLOTTEAUERBACH had been mingham, was invited and talked about his chromo- a research assistant to CREW; during thesecond world some-balance theory of quantitative inheritance. This war she discovered the first chemical mutagen, mus- asserted that + and - genes (those increasing and tard gas, and opened up wholea new field of research. decreasing the trait) are arranged in repulsion link- G. H. BEALEworked on Paramecium, and later de- ages. The net effect of a chromosome is minimal but veloped the new field of the genetics of the malarial it holds a large amount of hidden variation that can parasite. H. KACSER studied the genetics and enzy- be released by recombination. His theory was not well mology of metabolic pathways. B. WOOLFhelped received by the audience and he was criticized in a many people with his statistical advice, and was the forthright butinjudicious manner. MATHER,as a guest originator of the idea of realized heritability. W. G. speaker considerably senior to us, was understandably HILL’Swork has already been outlined. A later addi- Perspectives 141 tion was D. J. BOND,who worked on fungal develop- final illness. He died in 1989; his personality and ment. wisdom are greatly missed. From 1947 till 1955 there was a group of four, Did WADDINGTONkeep in touch with all the many funded by the Medical Research Council (MRC), staff and visitors? With the permanent staff he did, to working onthe mutagenic effects of radiation on some degree. We in the Unit seldom had any conver- mice. This was an outpostof the MRC’s Radiobiology sation with him, yet he often surprised us by knowing Research Unit at Harwell in Oxfordshire. Among its what we were doing and what we had found. He did members were T. C. CARTER,who later became di- this mainly, he said, by reading our papers. Of most rector of the ARC’S Poultry Research Center, andM. of the visitors, however, he knew very little. It was F. LYON,known for her work on X inactivation and not unknown for avisitor to have spent three years in the t locus in mice. Then, in 1958, the MRC set up the Institute and never to have spoken to him. Many anothergroup, the Mutagenesis Research Unit, of of the Ph.D. students came expecting towork under which AUERBACHwas director. It had five members WADDINGTON’Ssupervision. But often hewas away on among whom were B. M. CATTANACHand B. J. KILBY. his travels, and then the restof us had to come to the And finally, in 1963, the MRC founded the Epigenet- rescue, hastily thinkup suitable projects, and find ics Research Group under WADDINGTON’Sdirection space and facilities for them. with M. BIRNSTIELas his deputy. There were initially Although WADDINGTONseemed to take little day- four MRC staff in the group, but otherswith different to-day interest in what we did, he took great pride in funding worked in the group, among whom were K. the achievements of hisstaff. He was immensely proud W. JONES and, from theUniversity staff, J. 0. BISHOP, of the fact that in 1975 there were five Fellows of the R. M. CLAYTON,J. JACOB,A. JURAND and G. G. Royal Society among the Institute staff (WADDING- SELMAN.The group worked on development and TON, AUERBACH,BEALE, A. ROBERTSON,FALCONER) molecular genetics. with a sixth (MCLAREN) being elected shortly after WADDINGTONwas an inveterate traveler and was leaving. HILL was electedafter WADDINGTONhad widely known throughoutthe world. His interests died. This was out of a total of about 19 in the whole were wide-ranging, and not only in science. Among University and it elicited an article by a columnist in his 17 books was a lavishly illustrated one showing a newspaper (TAYLOR1975) in praise of the Institute. how modern art had been influenced by the ideas of Most organizations go through a cycle of growth science (WADDINGTON1969). WADDINGTON’Sbreadth and decline. The Institute was no exception. There of interests, and the reputation of the Institute, at- were three reasons why it grew the way it did: first, tracted many visiting research workers and Ph.D. and most important, WADDINGTON’Sdrive to expand; students, so that there were usually more visitors than second, the ready availability of funds at thetime; and indigenous staff. New people seemed to be arriving third, good fortune in the selection of staff. It reached almost every day and it was hard tokeep track of who the peak of its activities in the late 1960s and then was who and doingwhat. An idea of the numbers can started a gradual decline, for which I can see several be got from a list of people present in June, 1962. reasons. Possibly the prime reasonwas that WADDING- There were 22 permanent staff and 36 temporary TON’S interests moved from genetics and the Institute research workers. The visitors came from 13 different to futurology. In 1970 he went for two years to the counties in addition to the United Kingdom. Their Center for Theoretical Biology at Buffalo, and when fields of work arerecorded as: development (12), he returned heset up anew “School of the Man-Made quantitative genetics and animal breeding (lo), mu- Future” in the University. In consequence he was tation (7), Paramecium (4) Neurospora (2), and ga- seldom seen in the Institute from then till his death in metes (1). With so many people of such diverse inter- 1975. Fewer visitors came. Interest in most areas of ests the Institute was a lively and stimulating place. It research was shifting to the more molecular. Embryo was a great privilege to work there during that time. transfer and, later, theprospect of transgenic animals Naturally, lots of papers were published. In theperiod made classical quantitative genetics of less interest to 1961-1965 there were an average of 85 papers and animal breeding. Working in Edinburgh had been so 8 Ph.D. theses per year. attractivethat few staff membershad moved else- Meeting for coffee in the canteen was an important where. Consequently most of us were of an age when activity which kept staff and visitors in touch with each change becomes difficult. Funds were increasingly other. Some might say that this was a waste of research hard to get and,in the absence of new posts, very few time, but it was not; often we got valuable ideas and young people could be recruited to rejuvenate the advice. At first nearly everyone came, but later,as we Institute. got to know each other, the meetings broke up into To complete the history: before he went to Buffalo specialist groups. Of these, ALANROBERTSON’S coffee WADDINGTONwanted to shed his responsibilities in sessions became world famous and continueduntil his the Institute and in 1968 he resigned as Director of 142 D. Falconer the Unit. I was made Director in his place and at the WADDINGTON,C. H., 1939 An Introduction to ModernGenetics. same time was transferred to the University Depart- Allen & Unwin, London. WADDINGTON,C. H., 1969 Behind Appearance. Edinburgh Uni- ment in a personal chair; a year later I was made head versity Press, Edinburgh. of the Department andheld this position until J. R. S. FINCHAMcame to the Buchanan Chair in 1977. T. F. SOURCES C. MACKAYjoined the Department in 1980 when I Anonymous, 1950 Research in animal breeding and quantitative retired. ARC Units are normally terminated when genetics. A review of workin progress at the Institute of Animal Genetics, Edinburgh. Anim. Breed. Abstr. 18: 347-357. their directors retire.That ourunit survived a change CREW,F. A.E., 1971 The genealogy of the Poultry Research of director may have been due partly to its beginning Centre, Edinburgh. Br. Poult. Sci. 12: 289-295. as part of ABGRO and partly to the ARC’S difficulty FALCONER,D. S., 1977 Conrad Hal Waddington. Yearb. R. SOC. in finding new posts for its members.It did not, Edinb., pp. 58-60. HILL,W. G., 1990 Alan Robertson. Biogr. Mem. Fellows R. SOC. however, survive my retirement. By then most of its 36: 465-488. members had left or retired, and theUnit was finally HOGBEN,L., 1974 Francis Albert Eley Crew. Biogr. Mem. Fellows terminated in 1980. Very fortunately W. G. HILLwas R. SOC.20: 135-153. on the University staff and was not directly affected PUNNETT,R. C. (Editor), 1939 Procedings of the VII International Congress of Genetetics. Cambridge University Press, Cambridge. by the closure of the Unit. Anew cycle ofquantitative ROBERTSON,A,, 1977 Conrad Hal Waddington. Bibliogr. Mem. genetics has started under his leadership, with a new Fellows R. SOC.23: 575-622. and strong group. ROBERTSON,F.W., 1983 Genetics (in TwoHundred Years of the Biological Sciences in Scotland). Proc. R. SOC.Edinb. 84B 21 I- I am very grateful indeed to R. C. ROBERTSand W. G. HILLfor 229. reading the drafts and offering many helpful suggestions, and toB. WADDINGTON,C. H., 1973 Professor F.A. E. Crew. The Times, JAMIESONof the AFRC (formerly the ARC) for supplying some London, 2 June. dates. I ask forgiveness of any of my colleagues who find errors, WADDINGTON,C. H., and T. L. MASON,1951 Institute of Animal omissions, or misunderstandings. Genetics Edinburgh. Research 4: 3 15-3 19. UNPUBLISHED LITERATURECITED DEACON,MARGARET (undated) The Institute of Animal Genetics at Edinburgh-The First Twenty Years. LUSH,J. L., 1937 Animal BreedingPlans, Ed. 3. Iowa State College Institute of Animal Genetics, University of Edinburgh, Research Press, Ames, la. Reports. Five quinquennial reports for theyears 1947 to 1965. TAYLOR,W., 1975 More jolly good fellows; a Scotsman’s log. The MASON,I. L., 1950 The History and Workof the Institute of Scotsman, Edinburgh, 17 April. Animal Genetics, Edinburgh.