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Home , Birmingham, Brian Charlesworth, David Wilkie (surgeon), Grahame Bulfield, Jonathan Hodgkin, Lionel Penrose

Heredity7l (1993) 111—118 Genetical Society of

Genetics in the — The Last Half-Century

J. R. S. FINCHAM

Thisyear's 16th International Congress of in is the first to be held in the United Kingdom since the 7th () Congress in 1939, just before the outbreak of war. This historical essay is an attempt to chart the most important developments in U.K. genetics between these two Congresses or, more precisely, during the post-war period. I have tried to identify the institutions and schools that have had the greatest influence on the growth of our subject, and to trace the connections between them. I must to some extent be biased by my own partial view of the field, and I apologize for any serious omissions.

University, and establishing the fungus Aspergillus The early post-war scene nidulans as a model microbial eukaryote for genetic In1945therewere only a few centres in the U.K. for studies. The University Botany School was teaching and in genetics. There was the John another growth point for fungal genetics; Harold Innes Horticultural Institution (now the John Innes Whitehouse had already completed his Ph.D. on Institute), then located in Merton, South , with Neurospora sitophila and David Catcheside was about C.D. Darlington as Director. There were three institu- to import Neurospora crassa from CalTech. In R. A. tions concerned with plant breeding: the Plant Breed- Fisher's Genetics Department, L. L. Cavalli-Sforza was ing Institute in Cambridge and the Welsh and Scottish starting the work on the genetics of E. co/i that was to Plant Breeding Stations, near and Edin- lead to his discovery of one of the first two high- burgh, respectively. In University College, London, frequency-recombination strains, HfrC. The other, there was the Galton Laboratory, with L. S. Penrose as HfrH, was found a little later by W. (Bill) Hayes, of the the Galton Professor of , and the Department London Postgraduate Medical School in Hammer- of Biometry and Genetics with J. B. S. Haldane as smith. Hayes became the leader of the subsequent great Professor of Biometry. In Cambridge, R. A. Fisher expansion of bacterial genetics in the U.K. presided over a small Department of Genetics accommo- Two members of the John Innes Institution left to dated in an extension to his private house, while, on the seed genetics elsewhere. P. T. Thomas became Profes- other side of town in the Botany School, several young sor of Agricultural Botany in the University of , geneticists and cytologists were working under the Aberystwyth to teach a generation of Welsh plant guidance of D. G. Catcheside. In the Zoology cytologists and geneticists, several of whom (H. J. Department, E. B. Ford, with his population studies on Evans, H. Rees, E. A. Bevan, K. W Jones) are Lepidoptera, was a pioneer in ecological genetics. mentioned below. And in 1948 Kenneth Mather The immediate post-war period was a time of new moved to Birmingham University to found a new departures. In 1945, C. H. Waddington was appointed Department of Genetics and a powerful school of head of the Genetics Section of the newly established biometrical genetics, the most notable member of Animal Breeding and Genetics Research Organization which, from the mid-1950s, was John L. Jinks, who (ABGRO —laterABRO), funded by the Agricultural succeeded Mather to the Chair in 1965. In 1950 Research Council (ARC). IN 1946, the Organization Darlington and the John Innes moved from Merton to moved from London to Edinburgh, where Waddington a fine country estate in Bayfordbury, Hertfordshire. had been appointed to the Chair of Genetics in succes- sion to F. A. E. Crew. This appointment led to many MRC initiatives further developments, of which more below. Early On the other side of , Guido Pontecorvo Someof the most significant early post-war develop- was starting a new Department of Genetics at ments took place under the auspices of the Medical

111 112 J. R. S. FINCHAM

Research Council (MRC). Most important of all, in that Western General under the direction of W. H. it was about to revolutionize our subject, was the MRC Court Brown. Since 1969 it has been headed by H. Molecular Unit, established in Cambridge in JoIm Evans. Though modestly called a Unit, it became 1947 with Max Perutz as Director and John Kendrew a substantial Institute of international importance, with as his deputy. This Unit was concerned mainly with special strength in innovative technology, protein crystallography, but it also provided the venue characterization of human chromosome aberrations, for Jim Watson and 's momentous eluci- and research on radiation effects, with extensive use of dation of DNA structure. A few years later it played the mouse as a model mammal. host for a while to Seymour Benzer, who was then engaged on his demolition of the classical concept of the . The Cambridge MRC Unit was the direct Theexpansion of genetics in the Universities precursor of the MRC Laboratory of , established in 1962 in a large new building on The earliest Genetics Departments Hills Road with an all-star staff including Francis Crick, Atthe end of the 1940s there were independent Fred Sanger and Brenner. Crick and Brenner Departments of Genetics in four Universities —the led the molecular genetics group and attacked the long-established departments in Cambridge and Edin- coding problem, with results too well-known to need burgh and the new ones in Birmingham and Glasgow. description here. Edinburgh was the first of these to undergo major Another MRC initiative was their establishment of development. Soon after its move to Edinburgh in the Radiobiology Research Unit in Harwell. The inter- 1947, ABGRO was divided into the Animal Breeding ests of this Unit were by no means confined to radia- Research Organization (ABRO), a free-standing Insti- tion. In the 1950s, C. E. Ford carried out the first tute under H. P. Donald, later accommodated on the definitive studies on human in meiosis. University campus, and the Agricultural Research In 1961 Mary demonstrated X-chromosome Council ARC Unit of Animal Genetics, which was inactivation in female mice; she had started her work in assimilated into the University Department to form the Edinburgh where the Harwell Unit had an outpost in Institute of Animal Genetics with Waddington as Head. its early days. It was in Edinburgh that Charlotte This ARC Unit became famous for its work on the Auerbach had (during the war) discovered chemical theory and practice of selection, with Alan Robertson mutagenesis. In 1958 the MRC established a Muta- ( and cattle) and Douglas Falconer (mice) as genesis Unit there under her direction, which was the leading figures. This interest, combined with continued until her retirement. theoretical , continues to flourish in The 1950s saw the first flowering of bacterial and Edinburgh under the leadership of Robertson's student bacteriophage genetics in the U.K., and this too was and successor W. G. (Bill) Hill. It had, and still has, a largely funded by the MRC In 1957 an MRC Unit of strong teaching aspect in the form of a unique post- Molecular Genetics was set up under Bill Hayes at graduate course in animal breeding, run in collabora- Hammersmith Hospital. This soon became an tion with the ABRO staff. extremely strong international centre for the develop- With their emphasis on animals, random mating and ment of E. co/i genetics in all its aspects. At the same small populations, the Edinburgh quantitative geneti- time, Martin Pollock, at the MRC National Institute for cists differed from their Birmingham counterparts Medical Research (NIMR) at Mill Hill (North (who were also supported by the ARC) both in London), was working on bacterial enzyme induction, methodology and, to some extent, in . This interacting with 's group in Paris. difference at least partly accounts for our having two Pollock and Hayes and their respective colleagues general genetics journals, Heredity and Genetical (including almost the whole strong Hammersmith Research (the longer-established Journal of Genetics group) were later (1968) to join forces in a new, was taken by Haldane to India in 1957). Genetical impressively housed Molecular Biology Department in Research, which started in 1960, was edited from the the —asuccessful Institute of Animal Genetics by Eric Reeve, who has between an MRC Institute and a University teaching continued in the job to this day. department. Waddington's own interest included both selection In 1956, after the clarification by Tjio and Levan in and animal development. His experiments on selection Sweden of the human karyotype, the time was ripe for for response to temperature stress led to his concept of a new initiative in human cytogenetics, and the MRC genetic assimilation, and his thinking about develop- set up the Clinical and Population Cytogenetics Unit ment to his ideas about canalization —modelledas (now the Human Genetics Unit) at the Edinburgh epigenetic landscapes'. For support for these interests GENETICS IN THE UNITED KINGDOM 113 he turned to the MRC, and, from 1963, obtained fund- areas, notably cytology (Hubert Rees and later Gareth ing for an Epigenetics Unit, which obtained its own Jones) and bacterial genetics (Derek Smith, the Secre- building. The Edinburgh Epigenetics group, particu- tary of the present International Congress). larly Max Birnstiel, Ken Jones and John Bishop, made The Glasgow Department's great strength, derived a big mark in animal molecular biology, with from Pontecorvo, was always in Aspergillus genetics, pioneering work on such topics as rRNA , in situ which was important not only in itself but also for its hybridization of probes to chromosomes, and analysis impact on genetics in general. It resulted in the first of RNA transcripts. Most of the work on actual animal evidence (mainly arising from the work of Alan Roper development was done in the ARC Unit, especially by and R. H. (Bob) Pritchard) for the subdivisibility by Anne McLaren. There was never, in fact, any clear recombination of the functional gene. It also showed division of territory between the differently funded the potential, later much used in mammalian cell groups. Anne McLaren left in 1974 for University genetics, of parasexual systems of analysis, based on College, London, where Hans Gruneberg had been nuclear fusion and subsequent chromosome loss in working since the war on mouse developmental genet- vegetative cells. As we shall see, several of Pontecorvo's ics. younger colleagues of the 1 950s went on to become Another unique feature of the Edinburgh Institute of leaders of genetics in other Universities. The Asper- Animal Genetics was Geoffrey Beale's work on the gil/us tradition was maintained after John Pateman unusual genetic systems of Paramecium aurelia. In the succeeded to the Chair in 1970; the medical genetical 1970s his protozoan group extended its interests to connection was strengthened through Malcolm include pathogens such as trypanosomes (Andrew Tait, Ferguson-Smith. From the 1 960s, Glasgow University who has now moved to Glasgow) and malaria parasites also became a major centre for virus genetics; the lead- (David Walliker, who remains in Edinburgh). ing geneticist in the MRC Virus Research Unit was The Cambridge Genetics Department, distinguished John Subak-Sharpe, formerly a member of the Depart- but very small under Fisher, underwent a metamorpho- ment of Genetics, who became Director of the Unit in sis following the appointment of John M. Thoday to 1969. the Chair in 1959. Thoday, who obtained his Ph.D. under Catcheside in the Cambridge Botany School, Thebotanical connection had moved to University in 1948 and become Head of a new Department of Genetics there in 1954. Apartfrom the first four separate Genetics Depart- In Cambridge he obtained more adequate accommo- ments, the initial post-war expansion of University dation for the Genetics Department, first in the north- genetics was mainly in Departments of Botany. By ern outskirts of town but eventually (1976) in the 1950, two of David Catcheside's recent Ph.D. students, former Department of Agriculture building in the and myself, had left the Cambridge central area. During the 1960s the department Botany School for other Botany departments — developed all-round strength. Perhaps because of my Georgefor Trinity College, and I for . fungal bias, I give special importance to the Aspergillus In Cambridge, although Catcheside himself left in nidulans group (John Pateman, David Cove, Claudio 1952 for Adelaide, the genetical strength of the Botany Scazzocchio, Herbert Arst) who extended the range of School was maintained by Harold Whitehouse. In addi- the organism from formal genetics to biochemical (and tion to his own influential work on recombination, later molecular) genetics, setting in train a great expan- Whitehouse launched numerous Ph.D. graduates, sion of Aspergillus work internationally. This was the including Robin }lolliday and David Hopwood —and second subculture in the U.K. of Aspergillus nidulans their respective organisms, Ustilago and Streptomyces. from its Glasgow source, the first being in Sheffield, The role of P. T. Thomas in the Aberystwyth where J. Alan Roper, from Pontecorvo's department, Department of Agricultural Botany has already been succeeded Thoday as Professor and Head of Depart- mentioned. After his appointment, in 1966, as Director ment. In more recent years the Cambridge Department of the Welsh Plant Breeding Station he was succeeded has become the main U.K. centre for Drosophila in the University Chair by his former student Hubert genetics, mainly through the efforts of Michael Rees, who had spent some of the intervening years in Ashburner. Mather's department in Birmingham. Under Rees' In Birmingham, Mather's Department was always a guidance, Aberystwyth continues as a leading centre of stronghold of biometrical genetics, a position it main- plant cytology. tained under John Jinks. Like Edinburgh, but from a In 1957 Dan Lewis moved from the Headship of the somewhat different angle, it set up a taught post-grad- Genetics Department of the John Tunes to take the uate course in this field. But it also developed other Chair of Botany at University College, London, where 114 J. R. S. FINCHAM

he continued to pursue his interest in systems of self- Riley at the Plant Breeding Institute and was until incompatibility, now extended from cherries to the recently in the Cambridge Department of Genetics. fungus Coprinus lagopus. His main genetical colleague George Dawson, in Trinity College, Dublin*, got his there was David Wilkie (from Pontecorvo's group), Department in 1959, and stayed on to build it to its who became one of the pioneers of mitochondrial present strong position. And in 1950, the genetics of yeast. The Coprinus work was later taken University Department of Zoology (which already had over, to great effect, by Lewis's student Lorna Cassel- a senior population/ecological geneticist in Arthur J. ton, who is now a Professor in the Oxford Botany Cain) established a Sub-department of Genetics for Department. Philip M. Sheppard, who had come from E. B. Ford's Again in 1957, C. D. Darlington gave up the Direc- group in Oxford. This soon became a full Department, torship of the John Innes on his election to the Chair of with Sheppard in the Chair. He was a population genet- Botany in Oxford. This gave a botanical dimension to icist of great distinction, and it was a great blow when Oxford genetics, to complement E. B. Ford in the he died so prematurely in 1976. He was succeeded by Zoology Department. One of Darlington's first Donald Ritchie from Glasgow (and originally appointments was E. Alan Bevan, again from Ponte- from the Bill Hayes school), with a consequent shift of corvo's Glasgow group. Bevan later moved on to the emphasis in the department from population to bacter- Chair of Botany at Queen Mary College, London. His ial and viral genetics. investigation of the 'killer' phenomenon in yeast led to In 1966 both and started new his discovery of double-stranded RNA plasmids. Departments of Genetics. In Leeds, I was appointed to David Catcheside returned to the U.K. in 1957 (I the Chair, leaving the John Innes one year before its believe at Mather's instigation) to become Professor second move, this time from Bayfordbury to . and Head of Department of Microbiology in the Uni- I was followed in the Leeds Chair by David Cove in versity of Birmingham. This became in effect a depart- 1976. In Aberdeen the new Professor was H. John ment of microbial genetics. Catcheside and his Evans, who had been a colleague of C. E. Ford at the Birmingham students (including, most notably, Noreen Harwell MRC Unit. After his appointment in 1969 to Murray, now a Professor in Edinburgh) did definitive the Headship of the MRC Human Cytogenetics Unit in work on complementation maps and controls of Edinburgh, his place in Aberdeen was taken by Forbes recombination in Neurospora crassa. In 1964 he W. Robertson, previously in the quantitative group in returned to Australia to take charge of genetics in the Waddington's department. Research School of Biological in Canberra. The University of Wales established a Department of Genetics in in 1967, with the appointment The of John Beardmore (originally Thoday's student in spread of Genetics Departments Sheffield) to the Chair. and Newcastle- Fromthe mid-1950s, and especially in the 1960s and upon-Tyne followed in 1971 and 1972. The new early 1970s, the belief grew that any self-respecting Nottingham Professor, Bryan C. Clarke, originally a University should have a Department of Genetics. First student of A. J. Cain and Philip Sheppard in Oxford, in the field was Sheffield, as already noted. In the same brought a strong population/ecological interest to his year, 1954, my genetics laboratory in Leicester was Department, much needed at a time when the tide was given departmental status, but it was never much more flowing so strongly in the molecular direction. New- than a one-man show. When I moved in 1960 to castle appointed Stuart W. Glover, a bacterial geneticist succeed Dan Lewis in the John Innes Institute (joining originally from Dawson's Dublin department and after- Robin Holliday and Peter Day), the Leicester Genetics wards in the Hayes group, distinguished for his first Department fell into abeyance, a state from which it genetic analysis of restriction-modification systems. was rescued in 1963 by the appointment of Bob Those were the last separate University Depart- Pritchard, who built it to viable strength. Pritchard ments of Genetics to be created. Oxford never estab- came directly from Hayes' Molecular Genetics Unit in lished one, though in 1970, on E. B. Ford's retirement, Hammersmith and before that from Pontecorvo's it did start a new Genetics Laboratory within the department in Glasgow. Leicester now has one of our Department of . Fisher's old student strongest Genetics Departments, particularly famous Walter Bodmer, back from a long spell at Stanford for Alec Jeffrey's invention and development of DNA University, was the new Professor. At Oxford, Bodmer 'fingerprinting'. The current Head of Department is moved into human genetics, with an intensive investi- Gabriel A. Dover, who obtained his Ph.D. with Ralph gation of the HLA system. Two other Universities, Hull and (1973 and 1974), established Units of *Iam aware that I stray outside the United Kingdom at this point. Genetics associated with Botany and Zoology Depart- GENETICS IN THE UNITED KINGDOM 115

ments. Hull genetics, under David A. Jones, paralleled Glasgow and Dublin still maintain their separate status; that at Nottingham with its population and ecological the Department at Leicester, though part of a School of emphasis, while Manchester's specialities were Biological Sciences for teaching purposes, also retains mosquito genetics (R. A. Wood), cytology (R. D. Butler) a clear identity. and ecological genetics (L. M. Cook). Thegrowth of human and medical genetics Schoolsand mergers TheGalton Laboratory, which was founded as early as The1 960s was a time of establishment of completely 1904, has remained a leading centre for research in new Universities. The new foundations (Keele, East human genetics. The Annals of Human Genetics, for Anglia, Essex, Sussex, Lancaster, ) opted from the many years a premier journal in its field, was always start for integrated Departments or Schools of Biologi- edited from the Laboratory. The post-war Galton Pro- cal Science, which included genetics to different fessors of Human Genetics have been extents. The genetics group in the new University of up to 1965, Harry Harris between 1965 and 1976, and Sussex (1965) was by far the strongest: John Maynard then, until recently, Elizabeth Robson. Haldane was Smith and Brian Charlesworth (population/evolu- followed in the Chair of Biometry by C. A. B. Smith. tionary genetics), James H. Sang (Drosophila develop- Among the Laboratory's numerous important contri- ment), Neville Symonds and David Sherratt (bacterial butions, Harris's demonstration, with D. A. Hopkinson and bacteriophage genetics), and Bryn Bridges heading and others, of the large amount of enzyme polymorph- the MRC Cell Mutagenesis Unit. Charlesworth later ism in the human population, was a major landmark. moved to , and Sherratt to the Chair of Between 1962 and 1976 Harris was Director of the Genetics in Glasgow after John Pateman (following the MRC Human Biochemical Genetics Unit, first in the footsteps of Hayes and Catcheside) had left for University College Biochemistry Department and then Canberra. in the Galton Laboratory. When Harris left for Phila- In more recent years, several of the once separate delphia in 1976 the Unit was continued under D. A. University Departments of Genetics have become Hopkinson. merged in broader Departments or Schools. There Another distinguished London centre for human appear to be two main reasons for this —adesire for genetics, dating from 1946, was the MRC Blood and the perception that the way for- Group Unit, first at the Lister Institute and later in ward lies in interdisciplinarity. association with University College. This Unit will One consequence, perhaps paradoxical, is the split- always be associated with the names of its founding ting of the already interdisciplinary Edinburgh members R. R. (Rob) Race and Ruth Sanger. Race was Genetics Department between two Institutes, centred Director until 1963, and then Sanger took over until around cells/molecules/plants and animals/popula- her retirement in 1973. tions, respectively. It did make some sense, however, to During the later 19 SOs and 1960s there was a great unite the animal aspects of Edinburgh Genetics with increase in genetics and cytogenetics units in the hospi- Edinburgh Zoology, which had its own strong genetical tals. A major impetus was provided by improved tech- interests: in particular Aubrey Manning's behaviour niques for human karyotype analysis and the resulting genetics, Murdoch Mitchison's group and, in clinical applications. The big 1956 MRC initiative at the 1970s to 1980s, Peter Walker's MRC-funded the Edinburgh has already Mammalian Genome Unit. The Mitchison laboratory been mentioned. Before long, most of the larger hospi- launched the careers of some distinguished yeast genet- tals had cytogenetics laboratories. icists, including , and it was in Peter In several Universities, links were established Walker's Unit that Ed Southern invented his indispen- between clinical and academic departments. In Glas- sable blotting technique and (the current gow, the pioneer in clinical cytogenetics was M. A. holder of the Edinburgh Genetics Chair) pioneered the (Malcolm) Ferguson-Smith; he was, in the 1960s, a study of DNA methylation. Another Edinburgh zoolo- member of the University Genetics Department and, in gist, Linda Partridge, is about to take the Chair of 1974, became the Head of a new Department of Clini- Biometry in the Galton Laboratory at University cal Genetics, a position that he held until his move to College, London. Cambridge (University Pathology Department and Undergraduate courses leading to first degrees in Addenbrooke's Hospital) in 1989. In Birmingham the genetics have generally survived even where the leader was John H. Edwards, a population geneticist Departments no longer exist. At the time of writing, and paediatrician. He moved in 1961 from an M.R.C. Genetics Department at Cambridge, Leeds, Nottingham, Unit of Population Genetics in Oxford to the Bir- 116 J. R. S. FINCHAM mingham University Department of Social Medicine; Harris, not much concerned with plants. The Genetics in 1969 he became Professor of Human Genetics in the Department took on a rather fungal emphasis, with my University and Head of the Department of Clinical work on allelic complementation in Neurospora, Peter Genetics at the Birmingham Maternity Hospital. He Day's analysis (following Dan Lewis) of mating type in moved back to Oxford to succeed Walter Bodmer as Coprinus and, most notably, Robin Holliday 's Ustilago Head of the Genetics Laboratory in 1979. Oxford work which led to his path-breaking general model of already had strength in human/clinical genetics in the recombination. I claim some credit, with Brian person of David (now Sir David) Weatherall, a world Harrison, for resurrecting the work on genetic instabi- leader in the characterization of genetic haemoglobin lity (now known to be due to transposable elements) in deficiencies and Professor of Clinical Medicine. the snapdragon, Antirrhinum majus. This fascinating Over the last decade the leading hospital-associated system had been set aside since Kenneth Mather made laboratories have become more molecular in orienta- it the subject of his Ph.D. thesis in 1933. There was tion, adding the probing and analysis of DNA also a department of Potato Genetics, where Norman sequences to their previous cytological expertise. At St. Simmonds was engaged upon an enterprising project Mary's Hospital, London, for example, R. (Bob) designed to broaden the genetic base of the cultivated Williamson's group has been among the vanguard of potato by going back to the Andes. Jeffrey human gene cloners. carried on his masterly analyses of plant pigments, in An increasing role in human cytogenetic analysis has direct succession to Rose Scott-Moncrieff, whose pre- been taken by the Research Institutes, of which war work at Merton, prompted by Haldane, was one of the Imperial Cancer Research Fund (ICRF) Labora- the green shoots of biochemical genetics. tory in London, headed since 1979 by Walter Bodmer, of the Institute from its idyllic Bayford- must be the strongest. The ICRF has many advances in bury site to Norwich in 1967 was the consequence of a human cellular and molecular genetics to its credit, an Government Report that decreed that small Institutes outstanding example being the discovery by Peter doing fundamental research should be located in close Goodfellow (in collaboration with Robin Lovell- proximity to Universities. The new University of East Badge's group at NIMR) of the male-determining Anglia seized its chance. By the time of the move in element on the Y-chromosome. The ICRF has also 1967, the Director, Kenneth Dodds, had retired, undertaken research in basic molecular and cell genet- Harris had moved to a Chair in Oxford, Simmonds to ics, especially in relation to the control of cell division, direct the Scottish Plant Breeding Station, and I to a field to which Paul Nurse (the Genetical Society's Leeds to start the new University Genetics Department present President) has contributed notably. there. Peter Day and Robin Holliday had departed Peter Goodfellow has recently (1992) taken the some years previously, for Connecticut and the NIMR, Chair of Genetics in Cambridge. With the two adjacent respectively (Day returned in the 1980s to become Departments, Genetics and Pathology, headed respect- Director of the Plant Breeding Institute —thelast ively by Goodfellow and Ferguson-Smith, Cambridge Director as it turned out). The Jil was at a low ebb, but University is now exceptionally strong in human genet- under its new Director, Roy Markham the plant virolo- ics. With the human organism now virtually on a par gist, it rose on its new site like a phoenix from the with Drosophila as an object of intensive genetic study, ashes. Under the more recent Directorships of Harold we can expect a greater emphasis on human genetics in Woolhouse and Richard B. Flavell it has continued to University biology, both inside and outside the Medical gather strength. In the Genetics Department, David Schools. Hopwood and Keith Chater's group lead the world in Streptomyces genetics, and Enrico Coen and his ThePlant and Animal Institutes colleagues have used various transposable elements in Antirrhinum as gene tags to help open up a new field of Overthe post-war period, several plant and animal plant developmental genetics. research establishments outside Universities have been supported by the Agricultural Research Council (ARC —nowthe AFRC with the addition of Food to its ThePlant Breeding Institute/Cambridge Laboratory remit), and some of these have been in the forefront of ThePBI at Cambridge worked on a number of genetical research. different crop plants, but its most famous achievements were with wheat and barley. In wheat, Ralph Riley and TheJohn Innes Institute Cohn Law developed E. R. Sears' aneuploid analysis to a high art. This elegant wheat cytogenetics provided Inits later Bayfordbury days, the Jil included a strong the background for an extraordinarily successful Department under Henry (not Harry) breeding programme, due largely to John Bingham. GENETICS IN THE UNITED KINGDOM 117

PBI-bred varieties of both wheat and barley made on -specific genomic imprinting in mouse. In 1986, major impacts on the market. The Institute always ABRO, the PRC and Babraham (the first two amalga- combined practical breeding with basic plant science. mated on the PRC site at Roslin) were all placed under From the mid-1970s, Richard Flavell and his a single AFRC umbrella —theAFRC Institute for colleagues at the PBI were among the first to apply the Animal and Genetics Research. The new DNA technology to plants, with an initial concen- separation of 350 miles made joint administration diffi- tration on repetitive sequences and the cult, and the physiology and genetics are now once organizer. again separated under new names —theAFRC Babra- In the political climate of the 1 980s, and given the ham Institute and the AFRC (Edin- difficulty that the AFRC was having in funding all of its burgh), the latter headed by Grahame Bulfield. Institutes, the PBI's commercial success made it a The former ABRO building on the Edinburgh Uni- prime target for privatization. This is not the place to versity science campus is now occupied by a Univer- detail the somewhat confused circumstances surround- sity/AFRC Interdisciplinary Research Centre for ing the lucrative sale in 1987 of the breeding side of the Genome Research (CGR), under the leadership of PB! (not the basic science) to Unilever. Suffice it to say Richard Lathe and John Bishop. The CGR has a that, because of the status of the PBI as a Charity, the special interest in the potential of transgenic animals. lucre finished up, not in the hands of the AFRC or the With the Roslin Institute, the CGR and the University Treasury as everyone had expected, but rather with the Institute of Cell Animal and Population Biology (which Governors of the Institute. Now the Governors, includes the breeding and population interests of the deprived for the time being of AFRC support but care- former Institute of Animal Genetics), Edinburgh has a ful to retain AFRC goodwill, maintain the Cambridge position of dominance in U.K. animal genetics compar- Laboratory (as the non-privatized part of the PBI is able to that of Norwich in plant genetics. now called) in a well-equipped new building on the John Innes site in Norwich. Therise of molecular developmental genetics The move of the Cambridge Laboratory to the John Innes site followed shortly after the construction there Thelong-standing ambition of geneticists to be able to of the Sainsbury Laboratory, funded by the grocer and trace the connections between the genotype and the philanthropist David Sainsbury. This laboratory has, over the last decade or so, begun to be specializes in molecular plant pathology, and has a realized as never before. , of course, has been strong genetical aspect. With all three Institutions, Jil, molecular biology. The molecular analysis of develop- Cambridge and Sainsbury, working in liaison on the ment is being pursued in a number of University same site, and with significant strength in molecular departments of Zoology and Genetics, as well as in plant genetics in the nearby University of East Anglia, research establishments. In the forefront, just as it was Norwich may well be the strongest centre for geneti- in basic molecular genetics, is the Cambridge MRC cally oriented plant science in the world. Laboratory of Molecular Biology. One need only mention Peter Lawrence's work on compartments in Drosophila or Jonathan Hodgkin's on sex determina- AnimalInstitutes tion in Caenorhabditis. Some of the most original Onthe animal side, the institute most focused on mouse work has been at Babraham (, genetics was ABRO in Edinburgh, though the nearby imprinting), and the NIMR (Robb Krumlauf, Poultry Research Centre (PRC, founded in 1947) also clusters). The MRC Human Genetics Unit in Edin- did some genetical research. ABRO had close links burgh has also been increasingly involved in molecular with the ARC Unit of Animal Genetics in the Univer- studies of mouse development. sity until the Unit was discontinued on Douglas In Cambridge, the Cancer Research Campaign and Falconer's retirement in 1980. Some of ABRO's most the Welicome Trust have come together to establish a successful work was in pig breeding (John King) and new Institute of Cancer and . It selection theory (Charles Smith). In more recent years brings together, among others, and Ron A. John Clark has been making transgenic sheep, more Laskey from Cambridge Zoology, Martin Evans and with a view to pharmaceuticals than to improved meat Michael Akam from Cambridge Genetics, Azim or wool production. Surani from Babraham, and Anne McLaren from the The ARC/AFRC Institute for Animal Physiology at MRC Mammalian Development Unit in London. Babraham, near Cambridge, dates from before the war. The range of animals to be studied includes frogs, It acquired some genetical fame during the last decade mice, insects and no doubt others. This development or so, with the work of Azim Surani and his colleagues illustrates two features of the current scene: the 118 J. R. S. FINCHAM tendency for the strongest research groups to become evoked enthusiasm in some and boredom in others. concentrated in a few leading establishments, and the Molecular genetics had little apparent relevance to the abandonment of previous departmental boundaries. population/ecological wing, and vice versa. Those Can all this be claimed as part of our subject of whose expositions involved mathematics talked effect- Genetics? The answer is arguably yes, but, if so, we are ively only to each other. And even with the best will in laying claim to practically the whole of modern the world, it was becoming impossible to keep up with developmental biology. the expansion of knowledge in every area. From the 1960s, the Genetical Society started to Is include in each meeting both multiple concurrent ses- there a genetics community? sions of contributed papers and symposia, the idea of Whereas,in 1945, genetics was a minority interest the latter being to review special fields for non-special- among , viewed by most as specialized, unin- ized audiences. However, although the symposia were telligible or just irrelevant, it has now penetrated into usually excellent in their own terms, they were often, as every corner of biological science. Most biologists it turned out, directed mainly to the specialists. From would now recognize it as being at the root of every- the 1 960s, the population geneticists began to organize thing. This is a source of great satisfaction, but it also their own 'Pop Group' meetings, which became rather creates something of an identity crisis for geneticists. better attended than the Society meetings intended for What is Genetics? How do we distinguish ourselves the membership at large. Over the last few years, the from molecular biologists or biochemists, when half the Society has tended to abandon general sessions of con- papers in any leading molecular biological or tributed papers in favour of special symposia for biochemical journal are about identification of gene special interests. products and control of ? Another frank acknowledgement of diversity is the The other side of the geneticists' identity problem is publication of two Society journals: Heredity, a gift to the difficulty that people who are actually card-carry- the Society from C. D. Darlington in 1971, and Genes ing geneticists have in maintaining common interests and Development, founded on the profits from and a common pool of knowledge. Immediately after Heredity and started in conjunction with Cold Spring the war the British Genetical Society numbered only a Harbor in 1987. My guess is that hardly anybody con- hundred or so, and Society meetings consisted of tributing to the one journal would think of sending a common sessions of unsolicited contributions. Diverse paper to the other. as their interests were, the members managed to be So do we any longer have an identifiable subject? interested in most of what they heard. As the member- Perhaps we should stop worrying and adopt a prag- ship grew (its present strength is in excess of 1500) matic definition of a geneticist, perhaps as a person community of interest became more difficult to main- who thinks it worthwhile to subscribe to a Genetical tain. Bacterial genetics was a new world with a special Society. An alternative definition is someone prepared jargon and set of concepts that some of those out- to attend and even to help organize an International side the club found very difficult. Fungal genetics Congress of Genetics.