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GRAHAM PM Pm Int. J. Dev. Biol. 44: 51-55 (2000) UK mammalian development 51 Mammalian development in the UK (1950-1995) CHRIS GRAHAM* Zoology Department, Oxford University, Oxford, United Kingdom This is a brief history of mammalian developmental biology ing the Vietnam war and free child birth on the National Health research performed in the UK. It attempts to identify the wishes of system, Commonwealth citizens felt it a duty to visit relatives in the British society for developmental biology research, as expressed ‘home’ country, and it took longer for scientific funding to build up by politicians and refined by government funded research councils in continental Europe. and charitable agencies. The response of the mammalian develop- The first 20th century pulse is inextricably mixed with the distinct ment community of scientists is analyzed and found to have ferments of derivatives from the Marshall school of reproductive diverse origins. The major research achievements originated from physiology in Cambridge and Waddington’s Institute of Animal pure academic curiosity and very defined practical goals, with no Genetics in Edinburgh. Their programme was to improve farm obvious rules for making discoveries. During the period of this animal breeds and breeding, and this practical goal generated new history, the scientists became more effective at influencing the or neater techniques of super-ovulation (Edwards), embryo trans- wishes of British society. fer (McLaren), and persistent but failed attempts to produce Most British mammalian developmental biologists that have parthenogenetic mice and rabbits (Austin, Beatty, Braden, existed are alive, and they will have equally valid views about the Fischberg, Edwards, Pincus; see Beatty, 1957; Austin, 1961). This sequence of events and seminal influences. From the mid-1960s dual origin inextricably linked academic mammalian embryology on, the Society for the Study of Fertility and the British Society of with genetics, physiology and the application of science: a powerful Developmental Biology ensured annual contacts with workers in cocktail. North America and Europe, and it is impossible to isolate a uniquely In the 1945-1965 period, the public became acutely aware of the British contribution to the field. Instead, geographic location in dangers of nuclear weapons and the politicians’ sop was heavy Britain has set the limits to this economic, political, and scientific funding of mutation research, with the Medical Research Council history. Achievements in reproductive biology and pure develop- (MRC) maintaining six centres of radiation biology, of which only mental genetics are mentioned when they impacted on develop- MRC Harwell was to survive the cutbacks in the early 1970s. mental biology, but those subjects should write their own histories. Practical goals, when left to scientists, often lead to fundamental discovery. A measure of the embryology:genetics link was that the Stimulating funds and origins Mary Lyon hypothesis of X-chromosome inactivation depended on Mammalian developmental biology in Britain owes much to Abbreviations used in this paper: ARC, Agriculture Research Council; BBSRC, nationals of other countries who visited for longer or shorter periods Biotechnology and Biological Sciences Research Council; CRC, Cancer (hereinafter names in italic). This flux of talent could have been Research Campaign; EC, embryonal carcinoma; ES, embryonic stem; ICRF, promoted by the intellectual distinction of the UK. However, North Imperial Cancer Research Fund; MRC, Medical Research Council; NIMR, American scientists attributed at least equal importance to avoid- National Institute of Medical Research; SRY/Sry, Sex-reversed Y. *Address for reprints: Zoology Department, Oxford University, South Parks Road, Oxford OX1 3PS, United Kingdom. FAX: 01865 271228. e-mail: [email protected] 0214-6282/2000/$20.00 © UBC Press Printed in Spain www.ehu.es/ijdb 52 C. Graham events in the early embryo, and Bruce Cattanach’s observations on Mouse embryology paternal and maternal duplications gave the genetic seal to gene imprinting. An equally important consequence of radiation biology Cambridge contributed with the appointment of a self-effacing was the generation of mouse mutants, with the Medical Research professor with broad interests to the Darwin chair of Animal Council (MRC) Harwell and the Jackson Laboratory linking to Embryology. Bunny Austin fronted for two lecturers, Bob Edwards produce the free Mouse News Letter. The best descriptions of and Denis New, and sponsored a remarkable aggregation of mouse developmental anatomy were in the analyses of mutants by research students and postdocs, each beginning with a subject the genetics group of Hans Grüneberg at University College, apparently unrelated to mammalian development: Richard Gardner London, creating a tradition of meticulous anatomical description on sexing rabbit blastocysts, Martin Johnson on sperm antigens, which achieved monumental proportions (Kaufman, 1992). The Dave Whittingham on the metabolism of the preimplantation em- rich crop of mutants was decisive in nudging the British community bryo, and Azim Surani on mouse protein synthesis. Gillian Morriss- towards adopting mice as the “E. coli” of developmental biology, Kay in Anatomy carried forward Denis New’s work to analyse particularly because British developmental biologists were slow to teratology with cultured post-implantation stages. Each of these recognise the awesome power of Drosophila genetics. scientists was to head distinct mouse developmental biology Worries about radiation overlapped with and were soon sup- groups for at least 15 years. What sets these individuals apart from ported by fears of over-population: the new imperative was to others in the UK is that they were not diverted by talk of other model develop better methods of human birth control and to continue to systems, such as teratocarcinoma: they stuck closely to the improve farm stock productivity. For birth control, the scientists embryonic material and could thus analyse what really happened. were allowed to set the agenda and state that fundamental knowl- They also differed from their scientific offspring because they were edge was required about the metabolism of the mammalian able to pursue productive careers in the UK. conceptus. Money began to flow into Britain from the Ford Foun- Three fundamental features of mouse development were dation, World Population Council, and the Lalor Foundation to top exposed by this laboratory’s protégés. First in time, was cell fate up the increased funding from the Agricultural Research Council and cell commitment of the two cell populations of the blastocyst. (ARC, now BBSRC) and the MRC. The National Institutes of Health Second was the detailed investigation of the cell and molecular and the March of Dimes readily provided travelling fellowships for biology events which preceded the divergence of these two American citizens to postdoc in the UK and directly funded re- populations: arguably, this is the best study of the origins and search in the UK. At a guess, North American funds probably consequences of asymmetric cell division in any developing system. doubled the active mammalian development research community Third was the analysis of gene imprinting which has more to do with in the UK from 1960-1980 and the money mixed up research developmental genetics (later section). workers with different national and scientific backgrounds. The first study had the greatest impact on both developmental genetics and embryology because the techniques could be used to The prelude introduce single cells into the blastocyst for further development, and it thus provided one of the tools for the current gene manipulation The prelude to mammalian embryology was played by Anne techniques which inform about mouse development and much else McLaren, first defining the best conditions for embryo transfer and besides. then linking with Biggers to combine embryo culture with transfer Gardner and his colleagues brought their skills to Oxford, but (1958), thus opening up the pre-implantation window of opportunity their lineage analysis could not flower until there were good for embryo manipulation and subsequent development to term. markers. These were provided by somatic cell genetics. Somatic The final bars of the introduction were composed in the USA, where cell genetics required clear electrophoretic differences between fertilised 1-cell mouse embryos were grown to the blastocyst stage the isozymes of mouse and man, and one source of this variation and shown to be viable. The technique was quickly transferred by were the inbred strains of mice: isozyme variation between strains individual visits to the UK (Biggers, Brinster, Whitten; see Biggers, was to become the only cell marker for the first ten years of lineage 1981; Hogan et al., 1994). Everything was ready to launch an studies. During a one year postdoc in Anne McLaren’s laboratory attack on mouse development. in Edinburgh, Verne Chapman from Frank Ruddle’s group in the The increased funding and the atmosphere of the swinging 60’s States, visited most mouse development groups in the UK, created the freedom to just investigate the mouse embryo. It is odd demonstrated the techniques, and persuaded all to adopt isozyme that these possibilities were not noticed in the well funded centres markers. Combining clever and intricate micro manipulation with of learning, and it was an isolated academic environment which isozyme
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