The history of three scientific societies: the Society for the Study of Fertility (now the Society for Reproduction and Fertility) (Britain), the Socie ´te ´ Franc ¸aise pour l’E ´ tude de la Fertilite ´, and the Society for the Study of Reproduction (USA) John Clarke Department of Zoology, Oxford University, South Parks Road, Oxford OX1 3PS, UK Abstract Three scientific societies devoted to the study of reproduction were established in Britain, France and USA in the middle of the twentieth century by clinical, veterinary and agricultural scientists. The principal motivation for their establishment had been the study of sterility and fertility of people and livestock. There was also a wider perspective embracing other biologists interested in reproduction more generally. Knowledge disseminated through the societies’ scientific meetings and publications would bear upon human and animal population problems as well as basic reproductive physiology and its applications. New journals dealing with reproductive physiology, having worldwide appeal, were established in Britain and USA. The financial resources of at least one of the societies and its journal are directed towards charitable functions, including financial support for travel to scientific meetings, for visits to particular laboratories, and for research in the short term, including that of undergraduates. Perhaps the example of the British society has given rise to others having a more specialised focus, as well as to the formation of the European Society for the Study of Human Reproduction and Embryology. Ó 2007 Elsevier Ltd. All rights reserved. Keywords: Sterility; Fertility; Reproduction; Britain; France; USA 1369-8486/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.shpsc.2007.03.001 E-mail address: [email protected]Stud. Hist. Phil. Biol. & Biomed. Sci. 38 (2007) 340–357 www.elsevier.com/locate/shpsc Studies in History and Philosophy of Biological and Biomedical Sciences
The history of three scientific societies: theSociety for the Study of Fertility (now the Society
for Reproduction and Fertility) (Britain), the SocieteFrancaise pour l’Etude de la Fertilite, and theSociety for the Study of Reproduction (USA)
John Clarke
Department of Zoology, Oxford University, South Parks Road, Oxford OX1 3PS, UK
Abstract
Three scientific societies devoted to the study of reproduction were established in Britain, Franceand USA in the middle of the twentieth century by clinical, veterinary and agricultural scientists. Theprincipal motivation for their establishment had been the study of sterility and fertility of people andlivestock. There was also a wider perspective embracing other biologists interested in reproductionmore generally. Knowledge disseminated through the societies’ scientific meetings and publicationswould bear upon human and animal population problems as well as basic reproductive physiologyand its applications. New journals dealing with reproductive physiology, having worldwide appeal,were established in Britain and USA. The financial resources of at least one of the societies and itsjournal are directed towards charitable functions, including financial support for travel to scientificmeetings, for visits to particular laboratories, and for research in the short term, including that ofundergraduates. Perhaps the example of the British society has given rise to others having a morespecialised focus, as well as to the formation of the European Society for the Study of HumanReproduction and Embryology.� 2007 Elsevier Ltd. All rights reserved.
Keywords: Sterility; Fertility; Reproduction; Britain; France; USA
1369-8486/$ - see front matter � 2007 Elsevier Ltd. All rights reserved.
1. The general context for the study of reproductive physiology
The Society for the Study of Fertility (SSF), now the Society for Reproduction and Fer-tility (SRF), based in Britain, the Societe Francaise pour l’Etude de la Fertilite (SFEF),and the Society for the Study of Reproduction (SSR), based in the USA, focus most oftheir attention in practice on the biology of reproduction of Homo sapiens and other mam-malian species, some of them domestic or farm animals, but also common laboratory spe-cies as well as others of no obvious immediate commercial importance, but significant forthe study of comparative reproductive physiology. Nevertheless, other non-mammalianspecies, even invertebrates, sometimes feature in the meetings of one or other of these soci-eties and in their journals.
My research for this contribution began with a set of questions about the developmentof these now established scientific communities: what was the status of the study of repro-ductive physiology at and before the times these societies were formed? What were the cir-cumstances leading to the formation of these societies and, in the cases of SSF/SRF andthe SSR, their journals? Was there some overarching idea or problem that, in the 40s, 50s,and 60s of the last century, served to promote the study of reproductive physiology in, forthe most part, mammals? Did the problems formulated promote the development of new,particular techniques? Did these make possible the posing of new questions for reproduc-tive physiology? Here I draw on the literature generated by these societies and their leadingfigures, together with my personal experience as a member of SRF, in an endeavour to givesome answers, and to provide a history of research networks in reproduction from theirbeginnings to the present day.
An essential foundation for the investigation of reproduction, more particularly mam-mals, was created by F. H. A. Marshall (see Fig. 1) (Marshall, 1910).1 He provided in hisgreat book The physiology of reproduction not only the framework for the study of repro-ductive physiology but also some very important detail of the reproductive processes.There have been three further editions of this book, two of them greatly enlarged withmany contributors (Clarke, 2007), but still retaining essentially his original structure.Yet when in 1912 E. H. Starling’s2 renowned Principles of physiology was published, com-prising 1423 pages, just fifty-two of these were devoted to reproduction. Its further editionstreat reproduction at essentially the same length. What may have brought about this verylimited treatment of the physiology of reproduction in a book that has played such animportant part in medical education? Here are some possible explanations: (i) much ofthe contents are concerned with systems which determine the survival of individuals. Theseare naturally of immediate importance to clinicians, while reproduction is only (ironicallyspeaking) concerned with the survival of the species; (ii) reproductive physiology was amessy subject, and did lack the scientific precision of other parts of physiology that lentthemselves to rigorous formulation of hypotheses that could be tested experimentally;
1 F. H. A. Marshall, FRS (1878–1949) had done physiology at Cambridge in an undistinguished way, went onto the University of Edinburgh as a lecturer in physiology, returning to Cambridge to become Lecturer, thenReader, in Agricultural Physiology at the University’s School of Agriculture.
2 E. H. Starling, FRS (1866–1927) became Professor of Physiology, University College London. It is odd, in thepresent context, that it was he who, in 1905, coined the word hormone for a class of active agents one of whichW. M. Bayliss and he had identified as controlling the secretion of digestive enzymes by the exocrine cells of thepancreatic gland. Starling is perhaps better known for what is called Starling’s Law of the Heart, arising from hisinvestigations of the cardio-vascular system.
(iii) the study of reproduction was perhaps regarded as being slightly indecent, what AdeleClarke has referred to as the ‘illegitimacy’ of reproductive sciences (A. Clarke, 1998).
2. How did the Society for the Study of Fertility arise?
Before and after the publication of Marshall’s landmark book the results of researchin reproductive physiology were in Britain conveyed in the proceedings and publicationsof a wide range of scientific and clinical societies, such as the British Society for AnimalProduction, the Physiological Society, Royal Society, the Royal Society of Medicine, theSociety for Experimental Biology, the Society for Endocrinology, and correspondingorganisations in France, USA and elsewhere. However, the prospect of creating a scien-tific society dedicated exclusively to the study of reproductive physiology, whetherclinical or not, arose in the summer of 1944 when, in Exeter, Devon, four peoplewho, in the words of Arthur Walton3 ‘all had a common interest in problems concerned
3 Arthur Walton (1898–1959), agricultural scientist, later became Deputy Head of the Agricultural ResearchCouncil (ARC) Animal Research Station, Cambridge.
with reproductive physiology and the diagnosis, prevention and cure of impaired fertil-ity’, came together informally (Parkes, 1985). The participants were Margaret Jackson,4
Clare Harvey,5 Arthur Walton, and possibly G. I. M. Swyer.6 Annual, somewhat infor-mal meetings, under the auspices of the Family Planning Association, followed from this.The first of these was at the Animal Research Station in Cambridge in July 1945, at theinvitation of John Hammond.7 There were just seven participants: Margaret Jackson,Clare and Lesley Harvey,8 Arthur Walton, Alan Parkes9 and, as he wrote, ‘one ortwo others’.
By 1949, thirty or forty people attended the annual meeting (held in Edinburgh). This,as with the earlier ones, had a strong clinical component, with also some examination offertility and infertility in domestic animals. At that conference seven people set up theSociety for the Study of Fertility (SSF). The seven comprised five clinicians, together withArthur Walton, a senior researcher at the Animal Research Station, Cambridge, and AlanParkes, who was based at the National Institute for Medical Research, London. The inau-gural meeting of the SSF was held in 1950 at the Zoological Society of London, Alan Par-kes being elected the first Chairman. There were fifty members at the time of that meeting,and by 1997 this had increased to 1288, of which at least 354 were women. Besides theusual elected executive and other members of a learned society’s committee of manage-ment there were also seven ‘Trustees’ of the SSF, usually drawn from amongst thelong-standing members.
At the outset membership of the Society was formalised and required nomination by anexisting member of the founding group or of the Society as it grew. The applicant had (andhas) to provide evidence of her or his suitability for membership, which includes profes-sional qualifications. Election was by ballot of the membership of the Society (there aremuch more demanding requirements for membership of the Society for the Study ofReproduction in the USA, see below). When the management of the Society changed in2001 it was renamed the Society for Reproduction and Fertility (SRF), and two categoriesof membership were created: statutory and non-statutory members. Both types can attendscientific meetings of the Society, but there are small restrictions on the rights of each typerelating to voting and eligibility for grants from the Society.
4 Margaret Jackson (1898–1987), obstetrician and gynaecologist, Exeter and Devon District Hospital.5 Clare Harvey (1903–1996), Department of Zoology, University College of the South West, Exeter, later (1955)
University of Exeter.6 Formal records of the meeting do not exist. G. I. M. Swyer, FRCP (1917–1995), became Consultant in
Endocrinology, Department of Obstetrics and Gynaecology, University College Hospital, London.7 John Hammond, FRS (1889–1964) trained in natural sciences and agriculture at Cambridge. He was
renowned as an expert in practical problems of reproduction in farm animals and was appointed Lecturer, thenReader, in Agricultural Physiology, School of Agriculture, Cambridge University. He became Director of theAnimal Research Station, Cambridge (later the ARC Unit of Physiology and Biochemistry of Reproduction). Hecollaborated in some of his research with F. H. A. Marshall.
8 Leslie Harvey (1903–1986), Professor of Zoology, University College of the South West, Exeter, later ExeterUniversity.
9 A. S. Parkes, FRS (1900–1990) trained in agriculture at Cambridge, went on to do experimental biology atManchester University, University College, London University, and the National Institute for Medical Research,London, becoming there Head of the Division of Experimental Biology. In 1961 he returned to Cambridge as firstholder of the Mary Marshall and Arthur Walton Professorship of the Physiology of Reproduction.
The Proceedings of the annual meetings of the newly formed Society were published(at first by Heffers, Cambridge and then by Blackwells, Oxford) as Studies on Fertility,comprising Volumes 1–10, edited by R. G. Harrison, Professor of Anatomy, Universityof Liverpool. Through the initiative of Alan Parkes these gave way to the Journal of
Reproduction and Fertility, launched by him and a small number of senior members ofthe Society as Journal of Reproduction and Fertility Ltd., being a company of the Soci-ety, incorporated in 1960, ‘limited by guarantee and not having a share capital’ (Parkes,1985, pp. 332–334). It was thus legally and financially separate from, but otherwiselinked to, the Society and was its official journal. Parkes gave a number of reasons forthe establishment of the new journal: (i) many publications on reproduction had hithertobeen dispersed through existing biological literature; (ii) the mounting problem of theincrease in the world’s population demanded scientific studies for its control and a singlemedium was needed in which such investigations might be announced; (iii) the scientifictreatment of clinical problems, and research which might lead to increased productivityof farm animals (with favourable consequences for developed and less developed coun-tries), required a focussed outlet; (iv) the increased investigation of the reproductive biol-ogy of other species from or in the wild, which could give insight into their ecology,population dynamics and conservation. This called for a new journal consolidating theserelated problems.
A Council of Management for the journal (which in effect was the Editorial Board) wasdrawn from members of the SSF, and all members of the Society could become membersof the Company. At its outset the Council was chaired by Alan Parkes, and comprised ele-ven other members, all based in Britain, of whom seven were medically qualified,10 andthere were six corresponding editors drawn from six countries.11 The aims of the journalwere made clear in the first volume: ‘The Journal of Reproduction and Fertility publishesoriginal papers, abstracts of Proceedings and occasional reviews and bibliographies, deal-ing with the morphology, physiology, biochemistry and pathology of fertility’. The firstissue appeared in February 1960, edited by C. R. Austin, who had trained in VeterinaryMedicine at the University of Sydney and who, after some years as a research workerat the National Institute of Medical Research, was appointed to the Darwin Chair of Ani-mal Embryology at Cambridge University.
Prompted by the great increase in the number of publications dealing with problems ofreproduction, Parkes and Donn Casey (a graduate in agriculture from the University ofMelbourne, whom Parkes had by chance met in India in 1961) created the Bibliography
of Reproduction, comprising titles, authors and abstracts of papers dealing with reproduc-tion. Set up in Cambridge as ‘Reproduction Research Information Services Ltd.’ the firstnumber was published in 1963 and thereafter it appeared monthly (Parkes, 1985, pp. 88,352–354). Later David Cole and his small staff took over the Bibliography which had infor-mal links with SSF. Publication stopped in 1990 because of the increasing availability ofelectronic sources of abstracts.
10 E. C. Amoroso*, R. A. Beatty, P. F. M. Bishop*, J. L. Hancock, R. G. Harrison*, Margaret Jackson*, J. A.Laing, T. Mann*, L. E. Rowson, Linton Snaith*, G. I. M. Swyer* (*medically qualified).11 Australia (C. W. Emmens), Canada (R. L. Noble), France (C. Thibault, see n. 22 and Section 5, below), India
(V. Khanolkar), Israel (M. C. Shelesnyak), USA (W. O. Nelson).
Table 1Contents of three volumes of the Journal of Reproduction and Fertility and its successor Reproduction
Total number of pages and papers; other figures are percentages of total numbers.1 Bats, Hedgehog, Mole, Stoat.2 Armadillo, Murrah buffalo, Dusky leaf monkey, Hare, Spotted hyaena.3 Sea urchin.4 African elephant, Golden hamster, Grey squirrel, Tammar wallaby.5 Fish (Carp, Roach).6 Tape worm.
Table 1 presents an analysis of the contents of two volumes of the Journal of Reproduc-
tion (1960, 1980) and one volume of its successor, Reproduction (2005), giving the numberof pages and papers, the animals used in the investigations, and the laboratories and coun-tries in which the research was carried out. ‘Livestock’ include cattle, goats, horses, pigs,and sheep, and ‘laboratory animals’ embrace guinea pigs, hamsters, mice, rabbits and rats.For a discussion of the data, and a comparison with the official journal of the USA’s Soci-ety for the Study of Reproduction (Table 2), see Section 7, below.
Volume 1 (1960) was concerned with studies at a high level of physiological organisa-tion, such as pre-natal mortality; clinical problems of fertility; pregnancy block in micethrough the effect of alien males; oviducal transport of oocytes and embryos; ovarian tis-sue which had been frozen for storage and used later in grafting; the transfer of oocytesbetween individual rats; uterine activity; testis activity; testis blood supply; and the char-acteristics of semen and the passage of sperm through the male reproductive system. Therewere also studies of the survival of spermatozoa in the reproductive tract of more unusualmammals (bat, hedgehog, mole, stoat) and the reproductive physiological problems of a
Table 2Contents of three volumes of the Biology of Reproduction
Total number of pages and papers; other figures are percentages of total numbers.1 Deer mouse.2 Lizard, Fish.3 Bats, Cheetah, Deer mouse, Djungarian hamster, Ground squirrel, Musk shrew, Opossum, Squirrel monkey,
Vole, Zebra.4 Gold fish, Frog, Toad, Trout.5 Alpaca, Llama, Hyaena.6 White backed vulture.7 Alligator, Atlantic croaker, Flat head minnow, Medeka, Nile tilapia, Rice fields eel, Trout, Zebra fish.8 Crayfish.
bird species. The volume also carried the Proceedings of two meeting of SSF; unlike theJournal articles (see Table 1) these were mainly concerned with human clinical problems,the remainder focussing on common laboratory animals and livestock. As might beexpected from the composition of the Society’s early membership, more of the communi-cations were contributed from departments in medical schools or from hospitals than fromuniversity departments or research institutes. 85% of the communications came fromresearch groups in Britain, 10% from Ireland (where one of the SSF meetings had beenheld), and one each from Czechoslovakia, Sweden and USA.12
By the time of Volume 59 (1980), almost half way through the series to Volume 130(2005), the management of the journal had become more structured. There was an
12 Abstracts of papers given at the Society’s meetings continued to be published in the Journal until 1987 afterwhich they have made up a separate publication as an Abstract Series of the Journal.
Executive Editor, an Honorary Consultant Editor, an Executive Committee, Council ofManagement, and a Secretary. An analysis of the contents of the volume gives an indica-tion of some of the considerable transformations in reproductive physiology since thejournal began. There continued to be papers dealing with highly integrated systems, suchas genetic selection for litter size in sheep, or the influence of photoperiod on reproductionin sheep and hamsters. It had by then become possible to formulate questions moreprecisely and to take measurements with increasing specificity and precision. Moresophisticated methods of light and electron microscopy enabled finer and finer descriptionof organ structures in relation to their activity and, as a further example, thedevelopment of radio-immuno assay allowed more exact estimates in various physiologicalstates of hormone levels in tissue extracts or in body fluids. These procedures allowed thestudy of ovarian follicle dynamics, the experimental manipulation of follicular develop-ment, the effects of ovarian steroids on target tissues and feedback from them to theovaries.
But what is also notable in Volume 59 is the now routine use of in vitro methods for thepreservation of germ cells or embryos. Topics included investigation of embryo mortalityfollowing artificial insemination, the viability of frozen and thawed half embryos, the pro-duction of monozygotic twins by the separation of blastomeres in pre-implantationembryos and their transfer to the uterus leading to the production of live young. Relatedto such investigation are the analysis of epididymal and oviducal fluid, and influences onthem, and the survival of spermatozoa in media at different temperatures. Two serendip-itous advances made by Christopher Polge13 for the preservation of sperm (one arisingfrom the mislabelling of reagents in the lab, and the other through remaining in thepub longer than planned) played a very important part in these in vitro studies (Polge,1968; Parkes, 1985, pp. 147–151, 439). These investigations on germ cells and embryoswere basic to the later development of Artificial Reproductive Technology (ART). Manyof the publications in Volume 59 were strongly quantitative so that a common feature wasthe statistical analysis of results to establish whether the effects were likely to have beendue to treatments or to chance.
In 2001 the SSF and Journal of Reproduction and Fertility Ltd. were merged, and theSociety changed its name to Society for Reproduction and Fertility (SRF). At the sametime the Journal of Reproduction and Fertility became Reproduction, the official journalof the Society. Reproduction was formed from the merger of the Society’s two journals,the Journal of Reproduction and Fertility and Reviews of Reproduction. The reconstitutedSociety describes itself as ‘a registered charity, Limited by Guarantee, and under its Mem-orandum of Association must apply all its resources towards the enhancement of knowl-edge for the public benefit with reference to the reproductive processes and fertility in manand animals. As part of its charitable function it awards travel and other grants to indi-viduals to enable them to attend conferences and to undertake special projects’ (for exam-ple, to enable research to be continued between the end of one and the commencement ofanother longer term programme, or to support the laboratory work of undergraduates
13 E. J. C. Polge, FRS (1926–2006) trained in agriculture at Reading, began research work in 1948/49 under AlanParkes at the National Institute for Medical Research, London, and became Director of the ARC Unit of thePhysiology and Biochemistry of Reproduction, Cambridge, which became incorporated into the AfRC Instituteof Animal Physiology, Babraham near Cambridge, while still retaining its identity at Huntingdon Road,Cambridge. See Polge (2007).
carrying out investigations with an established research group).14 This particular function,which had been long standing, had been formalised by Alan Parkes in the 1960s.15 Theaffairs of SRF are in the hands of a Council made up of the usual office bearers of scientificsocieties, and four Advisory sub-Committees. Three of these are quite usual (Finance,Publications, Meetings) but the fourth (Education) is a notable recognition of that partof the stated aims of the Society that refers to ‘knowledge for the public benefit’.
The journal in its present guise (Volume 130, 2005) has an Editor-in-Chief, a reviews orsometimes a US editor, an Editorial Board for reviews of sixteen people, drawn from Aus-tralia, Britain, France, New Zealand, and USA, and an Editorial Board for researchpapers of forty people (the members being from Australia, Belgium, Britain, Canada,France, Ireland, Japan, New Zealand, Spain and USA). This contrasts with the organisa-tional structure adopted for Biology of Reproduction in the USA, whose administration isdrawn exclusively from scientists in North America (see below). Table 1 gives an analysisof the contents and authors of this recent volume. Although the research reported there ispredominantly focussed on events at the cellular and sub-cellular level, some attention hasbeen paid to processes in more highly integrated systems. This includes comparativeaspects of reproduction and also clinical problems. Thus, there are papers on wallabies,elephants and fish. On the clinical side there are reports on the fertility of women; the char-acteristics of developing embryos; the influence of leptins in maternal adipose tissue; andthe control of labour.
Investigations at a cellular or sub-cellular level have focussed on germ cells and earlyembryos (including ionic exchange and structural changes before and after fertilisation).Other papers deal with genes involved in hormone action, and their expression in humanovarian follicles, oocytes and embryos. There is now understanding of control by genes ofsome of these processes: the role of the genome in the regulation of reproductive processeshas been greatly helped by the development of strains of mice which have had their geneticmake up deliberately altered (‘knock-out’ mice). These insights have included betterunderstanding of genomic imprinting (Swales & Spears, 2005). Reports of in vitro meth-ods using sheep and cattle embryos contribute important information on fecundity, fertil-ity, and embryonic development (for example, the interference of a virus in development).Other investigations reported include the transfer of somatic cell nuclei into enucleateoocytes, in one case leading to in vitro development of horse embryos, later yieldinghealthy offspring. Overall the contents admirably match the stated scope of the journal:‘Reproduction publishes original research articles and topical reviews on the subject ofreproductive biology. Its focus is on cellular and molecular biology of reproduction,including the development of gametes and early embryos, assisted reproductive technolo-gies in model systems and in a clinical environment, reproductive endocrinology andreproductive physiology. Emerging topics including cloning, the biology of embryonicstem cells and epigenetic effects on embryonic development are encouraged’.16
14 See Reproduction, 130(1) (2005), p. ii.15 Parkes wrote: ‘By the time I retired, the Journal had started on its prosperous phase and later set up a grant
fund for the assistance of members of the society : : : to facilitate matters I offered to act as Settlor by contributing£100 to set up the Trust for Education and Research in the Biology of Reproduction’, whose main capital thencame from the Journal of Reproduction and Fertility (Parkes, 1985, p. 337). Parkes retired from his Professorshipat Cambridge University in 1967.16 See Reproduction, 130(1) (2005), p. ii.
Although much of the work published in the Journal of Reproduction and Fertility andin Reproduction has been done on conventional laboratory or domestic animals and live-stock, comparative studies have also been an integral part of the life of the Society. Pastnumbers of the journals include (among others) studies on the reproductive physiology ofthe Rhesus Monkey, Black Bear, the Indian Rhinoceros (albeit in a zoo), the Grey Seal, abat, a Grasshopper Mouse, the (wild) White-footed Mouse of North America, the PersianSturgeon, the Trout and the Spiny Dogfish. The comparative viewpoint was promoted in afocussed way through the creation by Alan Parkes of ‘Symposia on the Comparative Biol-ogy of Reproduction’, linked to the SSF. The first of these, entitled ‘Comparative Biologyof Reproduction in Mammals’, was held in London in 1964 and considered problems inthe reproduction of a very wide range of mammals from Primates to Marsupials, with con-tributors from Australia, Britain, East Africa, France, India and USA. The second Sym-posium, ‘Biology of Reproduction in Mammals’, was held in Nairobi in 1968. There wasan introductory section on general morphological and on physiological aspects of repro-duction, but the rest of the Symposium dealt largely with a very wide range of problems inAfrican mammals, based often on animals in the wild, though some mammals from otherparts of the world were also considered. Topics ranged from Baboon menstrual cycles andtheir pre-implantation embryos, to the reproduction of the Right Whale in Southern Afri-can waters. Other species of primate, and many species of African grazing animals androdents were also discussed. The contributors came from Biafra, Britain, Canada, France,Germany, Kenya, the Netherlands, South Africa, Tanzania, Uganda, USA, Western Cam-eroon and Zambia.
The third Symposium, held in Edinburgh in 1972, was entitled ‘The Environment andReproduction in Mammals and Birds’. Consideration was given to the roles in reproduc-tion of photoperiod, ambient temperature, nutrition, rainfall, pollution, olfaction, popu-lation density, human activity and animal populations, as well as discussing humanreproduction.17 The final one was held at Canberra in 1976 under the auspices of the Aus-tralian Academy of Sciences, the Australian Society for Reproductive Biology and theSSF. It dealt with ‘Reproduction and Evolution’ as illustrated by mating systems, sexualselection, season of parturition, interspecies fertility, the evolution of sex determination,the comparative reproduction of male vertebrates, the ovarian cycle and luteal function,embryo-maternal interactions, and the evolution of viviparity.18 These Symposia demon-strate vividly Parkes’s commitment to a comparative approach to reproduction, bringingwith it a corresponding global array of organisms and research workers.
4. Scientific meetings of the Society
Until very recently two scientific meetings, based on University departments, were heldannually, made up of symposia with invited speakers, as well as short oral and postercommunications by members of the Society. The Winter meeting was in London and aSummer one elsewhere in Britain or Ireland. However, now there is just one meeting, inthe spring or summer. Meetings have often been held jointly with one of a number of dif-ferent scientific societies having shared interests with SSF/SRF, for example, the British
17 Contributors were from Australia, Britain, Canada, Germany, Hong Kong, Kenya, Netherlands, Poland, S.Africa, Sudan, Sweden, USA, and Zambia.18 Contributors came from Britain, Canada, Israel, New Zealand, S. Africa, USA, and Australia.
Neuroendocrine Group, the Reproduction, Growth and Development Group within theNutrition Society, the British Society for Developmental Biology, the British AndrologySociety (BAS), the British Fertility Society (BFS), and the Society for the Study of AnimalBreeding.
There have also been a number of meetings with equivalent scientific societies on themainland of Europe—starting (March, 1975) with a meeting in Tours with the Societe
Nationale pour l’Etude de la Sterilite et de la Fecondite, as it was originally called, laterbecoming the Societe Francaise pour l’Etude de la Fertilite (SFEF, see below). This liaisonwith the French society was possibly seeded by the tradition, established by ThaddeusMann,19 of a biennial meeting at the laboratories of the Institut National RechercheAgronomique (INRA), Jouy-en-Josas, near Paris, between French reproductive physiolo-gists and those from the Agricultural Research Council (ARC) Unit of Physiology andBiochemistry of Reproduction, Cambridge. There have been a number of furtherFranco-British meetings as well as those with other scientific societies in Europe whichare concerned with reproductive physiology—German, Dutch, and Flemish societies.The coming together of the SSF/SRF with related societies has been very fruitful scientif-ically and most agreeable socially.
A number of special named lectures have been given annually at meetings of the SSF/SRF to honour those who have made notable contributions to the Society’s organisationor its proceedings: a Parkes lecture, honouring Alan Parkes’s crucial role in the establish-ment of the Society, quite apart from his important contribution to the understanding ofreproductive physiology; an Amoroso lecture, in memory of E. C. Amoroso,20 not only forhis scientific research but also as a ubiquitous participant in the Society’s meetings and inthe management of the Journal; a Hammond lecture, an acknowledgment of the veryimportant contributions made by John Hammond to the understanding of breeding indomesticated animals; an (Arthur) Walpole lecture, supported by a fund created by hisImperial Chemistry Industry colleagues, partly in recognition of his work which bore uponthe development of the contraceptive pill. These lectures now appear to have been replacedby an annual Distinguished Scientist Lecture, and a Patrick Steptoe Memorial Lecture.21
In addition to commemoration by lectures, the SSF/SRF has awarded annually a Mar-shall Medal (in memory of F. H. A. Marshall) to a scientist who has contributed signifi-cantly to the understanding of animal reproduction. Alan Parkes was responsible for thecreation of the medal, which is usually presented at the dinner of the Society held duringthe annual Spring or Summer scientific meeting. The award is not confined to members ofthe Society. There have been forty-one recipients of the Marshall Medal. Of these, threehad made fundamental contributions to the understanding of endocrinological aspects
19 Thaddeus Mann, FRS (1908–1993), biochemist, came from Poland in 1935 to work at the Molteno Institute,Cambridge University, having been awarded a Rockefeller Fellowship. He became Director of the ARC Unit ofPhysiology and Biochemistry of Reproduction, Cambridge. He never returned to Poland, but his origin on themainland of Europe may have encouraged him to form Continental connections.20 E. C. Amoroso, FRS (‘Amo’) (1901–1982) was a West Indian from Trinidad who qualified in medicine in
Dublin, and later did research in Freiburg, Berlin, and at University College London, before becoming Professorof Physiology at the Royal Veterinary College, London.21 Patrick Steptoe, FRS (1913–1988) was Consultant Obstetrician, Oldham and District General Hospital,
Lancashire. He and R. G. Edwards, FRS, of the Department of Physiology, Cambridge University and laterBourn Hall, Cambridge, were responsible for the production by in vitro methods, and birth of, Louise Brown, thefirst such child. See also n. 29 below.
of reproduction; nine were trained in medicine, of whom seven did research arising from,and contributed to the solution of, clinical problems, often using sheep as experimentalanimals, while two investigated the epidemiology of human fertility; a further eleven fromuniversity departments of animal or agricultural science or similarly orientated researchinstitutes, tackled problems bearing on livestock production and eighteen from universitybiology departments or biology research institutes, had made contributions of importancefor the understanding of the reproductive physiology for a broader range of animals. Thefirst and third (as well as several later) were from USA (1964, C. G. Hartman; 1967, H. M.Evans), and the second (1966) had initially carried out research in Germany (B. Zondek).None had been members of the Society at the time of being presented with the medal.Charles Thibault, from France, who in addition to his important research contributedin a crucial way to the formation and development of the Societe Francaise pour l’Etudede la Fertilite (see below), was also awarded the Medal in 1980.22 There are only fourwomen in the list, which surely indicates a surprising sex bias. Two of them were medicallyqualified, and two non-medical experimental biologists. The first was Hilda Bruce,awarded the medal in 1974, followed by Margaret Jackson (see page 343) in 1979, AnneMcLaren in 1985 and Hannah Peters in 1992. A somewhat similar biased and puzzlingsex ratio can be seen in the various awards made by the Society for the Study of Repro-duction (USA).
5. Formation of the Societe Nationale pour l’Etude de la Sterilite et de la Fecondite renamedby 1984 the Societe Francaise pour l’Etude de la Fertilite
In 1954 professors Paul Funck-Brentano and Raoul Palmer, who were engaged inParis23 with other endocrinologists, gynaecologists, and obstetricians in the treatment ofclinical problems of human reproduction, proposed the formation of a Societe Nationalepour l’Etude de la Sterilite et de la Fecondite. The executive was set up in 1955 with Pro-fessor Funck-Brentano its first President. The founders wanted to incorporate not onlythose research workers tackling problems of fertility in humans but also others concernedwith domestic animals and livestock, to form a learned society comprising gynaecologists,obstetricians, veterinarians and biologists. By 1955 there were 240 members, includingCharles Thibault, and apparently, even then, clinicians from Italy, Morocco, Mexico, Par-aguay, Sweden, Syria and Vietnam.24 Thibault promoted a more comprehensive biologicalapproach to reproduction extending beyond the interests of clinicians. This may to someextent have stemmed from his broad biological training. He shared some personality attri-butes with Alan Parkes. Their careers in a way came together, though they had rather dif-ferent starting points (Poupardin, 2002).
22 C. Thibault (1919–2003) trained as a biologist, whose doctorate comprised two quite different problems: onewas concerned with colour vision in fish, and the other with parthenogenesis in mammals. He did this work whilea member of the Centre National de Recherche Scientifique (CNRS). He then became an Assistant in Biology atthe University of Paris, after which, as a research worker with the Institut National de la Recherche Agronomique(INRA), he became increasingly engaged with the problems of reproductive physiology with a strong emphasis onlivestock. He became both Director of INRA and of its Department of Physiology, was later appointed Professorof the Physiology of Reproduction, University of Paris VI, and then President of the CNRS.23 This work was associated with Hopital Broca, Assistance Publique des Hopitaux de Paris.24 Vietnam’s participation may be explained by links established during the period of French colonialism in
The first colloquium of the Society, held in 1955, was concerned with the oviduct ofwomen in health and disease. Between 100 and 150 people participated in that meeting.There have been (up to 2004) a further thirty-eight colloquia, embracing a wide rangeof reproductive problems. Between 1969 and 1972 and thereafter, the clinicians werejoined by those working on the physiology of reproduction of domestic animals, livestockor other animals. Many of these research workers were from laboratories of the CentreNational de la Recherche Scientifique (CNRS) and the Institut National de la RechercheAgronomique (INRA). Along with clinicians, they contribute research results to the Soci-ety’s proceedings and serve as ‘Rapporteurs’ for the colloquia. These have dealt withreproductive physiology from clinical and non-clinical points of view, embracing the epi-demiology of fertility and sterility, investigations at the level of whole animals and theirintact reproductive organs, and the genetic, neurophysiological, endocrinological, immu-nological, cellular and sub-cellular aspects of reproduction. This included, of course,gametogenesis, fertilisation, embryo implantation, pregnancy, and parturition. The sym-posia held between 2002 and 2004 dealt with ‘Age et reproduction’, ‘Dysimmunite etreproduction’, and with assisted fertility, focussing on ‘Insemination Intra-uterine: Quand,Comment et pour qui, Pourquoi?’ (Reynaud, 2005). The broad approach to reproductivephysiology has sustained the initial objectives of the Society. There appears to be no offi-cial journal for the SFEF. The publications of its members are spread through existingclinical and biological journals.
6. Formation of the Society for the Study of Reproduction (SSR)
The origins of the Society for the Study of Reproduction in the USA can be traced to1966 and the efforts of Philip Dziuk, at that time Assistant Professor at the University ofIllinois. He wrote to persons he knew who were doing research on problems of reproduc-tion, suggesting that there perhaps should be formed a society, structured after the modelof the Society for the Study of Fertility ‘established in England in the late 1950s and man-aged to amalgamate diverse interests and groups’. It is perhaps significant that just beforethis Dziuk had been working for some months in Chris Polge’s laboratory in Cambridge,for Chris was already a dedicated member of the SSF. At a Symposium on AnimalReproduction held in June 1967 at the University of Illinois, Champaign, and sponsoredby the American Society of Animal Science, Dziuk proposed the formation of whatbecame the SSR. In the ensuing discussion it was apparent that ‘not all in attendancewere in favour of establishing a new society. Many of the established investigators viewedthe new group as divisive to existing groups rather than unifying’ (Dziuk, 1993). Never-theless the Society was organized formally in June 1967 and R. M. Melampy, Depart-ment of Animal Science, Iowa State University, Ames, was appointed President. Theaims of SSR were ‘to provide a forum in the form of annual meetings and eventuallya journal for all research workers interested in reproduction : : : regardless of whethertheir research interests are with wild animals, laboratory species, domestic animals orman’. Hitherto, there had not been such a common meeting ground for the medical pro-fession, animal scientists, zoologists and all others concerned with problems of reproduc-tion (Nalbandov, 1968).
The first of what became annual scientific meetings of the SSR was held in 1968 at Van-derbilt University, Nashville, Tennessee. 213 members of the Society participated as well
as 43 students and 56 non-members, and there were 85 abstracts of papers. In 1969 themembership was 740 (Dziuk, 1993). By 1969 management of the SSR had become morehighly structured.25 J. D. Biggers became President. He worked in the Laboratory ofHuman Reproduction and Reproductive Biology, Harvard Medical School, had trainedat the Royal Veterinary College, London, and had done research in Britain and Australiabefore going to USA. Besides the President, there were a number of executive officers aswell as an Executive Director, nine Directors, and two trainee representatives. There arefour types of members of the SSR: Regular, Associate, Sustaining and Trainee, and therequirements for membership are quite rigorously specified.26 To become a Regular mem-ber, candidates are required to have a doctorate or equivalent qualification, and to submita bibliography of their published papers in the area of reproduction. They must have aminimum of three publications with the applicant as the first or last author, or five pub-lications with the applicant as the first or last author of at least one. Applicants who citepublications of theirs in a language other than English must submit one reprint of eacharticle for proper evaluation. Associate members may be drawn from research associatesor senior technicians engaged in the area of reproduction, or those who may be interestedin the science of reproductive biology but not actively engaged in it, or others who areengaged in full-time clinical practice but have not published, or do not expect to publish,in the area of reproduction. Sustaining members are individuals or firms connected to, orinterested in, the advancement of knowledge about human and animal reproduction. Trai-nee members are students training in reproductive biology and are recommended by a reg-ular member. That the Society has flourished is clear from its membership (in 2004) of1790, while the annual meeting of that year attracted 1288 participants and 824 abstracts(Hoyer, 2005).27
The SSR gives a number of annual awards to those who have contributed in a particu-larly important manner to the study of reproduction. As with the recipients of the MarshallMedal from the Society for Reproduction and Fertility (see Section 4), so with SSR there isa striking inequality in the sex ratio of the awardees. Up to 2005, 2 out of 37 Carl Hartmanawards, 4 out of 29 SSR research awards, and 3 out of 21 Distinguished Service awardshave been to women. Further research could perhaps establish whether these ratios faith-fully represent the contribution that women have made to the study of reproduction.28
7. A new journal: Biology of Reproduction
In 1969 the SSR started a new journal entitled Biology of Reproduction. Its first Editorwas H. H. Cole of the University of California, Davis, with six Associate Editors and anEditorial Board of forty-six scientists. After Volume 1, the journal greatly increased in size(see Table 2), but staffing for Volume 36 (1987) was essentially the same as for Volume 1.Although Volume 73 (2005) is no bigger than Volume 36, the administration of the journalhas become much larger. By December 2005 there were two Co-Editors-in-Chief, ten or ele-ven Associate Editors, a Board of fifty-one Reviewing Editors, plus a Consulting Editor, aManaging Editor, a Deputy Managing Editor and an Editorial Assistant. The executive of
25 Biology of Reproduction, 1 (1969).26 See Biology of Reproduction, 73(6), (2005).27 Professor Patricia Hoyer, Department of Physiology, University of Arizona, Tucson, USA.28 See, for example, Biology of Reproduction, 73(2, 3 or 4), (2005).
the journal has introduced a number of small innovations, which make it easier for readersto find those parts of an issue that are of particular interest. In the table of contents of anissue, each paper is accompanied by a one or two line summary of its objectives. There isalso a ‘Contents category’ index embracing gamete biology, mechanism of hormone action,ovary, pituitary, pregnancy, testis, and reproductive technology. In a short additional sec-tion, entitled ‘Biology of reproduction highlights’, the Editors draw attention to particu-larly noteworthy research. Publications in the Biology of Reproduction, and no doubtcommunications at meetings of SSR, cover the same range of problems and techniquesas is embraced by the Journal of Reproduction and Fertility and by Reproduction, namelyfrom whole animals down through various layers of physiological organisation to the cel-lular and sub-cellular level. Research on problems of reproductive biology in humans isquite frequently reported, though this may not be directly on clinical cases but on the exam-ination of tissues, hormone titres and other parameters derived from clinical material.There are also investigations on more unusual or exotic animals (see Table 2).
Table 2 presents an analysis of the contents of three volumes of the journal (1969, 1987,2005) adopting the categories used in Table 1 for comparison. Simple perusal of Tables 1and 2 indicates that the distributions for the two journals of types of animals used and theplaces (laboratories and countries) where research was carried out are essentially the same.It is not surprising that contributions to the first volume of each journal were from labo-ratories in the country of origin of the journals, but that each journal later developed aninternational appeal. Nor is it surprising that the size of each journal and the number ofpapers published has increased very considerably. There are perhaps two facts upon whichto comment: the appreciably higher incidence of studies on humans in the first volume ofthe Journal of Reproduction and Fertility than in Volume 1 of the Biology of Reproduction,which could be regarded as consistent with the human focus at the creation of the Britishas compared with the American society; and the wide range of exotic animals on whichstudies have been published in both journals.29 There are sometimes quite large numericaldifferences between the British and American journals both in the contributions to themfrom hospitals or medical schools, university departments and research institutes, as wellas in the countries from which they originated; however, the many and varied factorswhich produced these figures preclude any simple interpretation.
8. Concluding remarks
We have seen that the number of members of the SSF/SRF has risen from about twentyor thirty to about 1288 in 1997, but there has been a decline since then to a little over 600.Membership, at least in its peak period, was drawn from many parts of the world. How-ever, with increasing specialisation in the nature of the problems being investigated, newsocieties have developed: at least that is true of what has happened in Britain. The SSF canperhaps be regarded as the parent society out of which grew in 1972 the British FertilitySociety (BFS) and in 1977 the British Andrology Society (BAS). Contributing to the for-mation of the BFS was ‘dissatisfaction with the Society for the Study of Fertility, whichappeared dominated by animal scientists with minimal human concerns’ (Reiss, 1997).
29 This fits in both with Alan Parkes’s broad intentions for the English journal and his Symposia for thecomparative biology of reproduction, and with the stated aims of the American society.
While this may have reflected the general orientation of the Society after its early years, theSSF/SRF and its members have continued to make notable contributions to the develop-ment of clinical medicine. For example, R. G. Edwards’s30 experimental studies on spermand oocytes in laboratory animals ultimately led, in collaboration with Patrick Steptoe, tothe birth of the first baby conceived by IVF techniques they had painstakingly andpatiently developed (Steptoe & Edwards, 1978). There is also the example of the experi-mental investigations by David Baird31 and his colleagues on sheep reproductive physiol-ogy. He has remarked: ‘the SSF, French Society and ASRB (Australian Society forReproductive Biology) in particular, have made a major contribution to reproductive sci-ence and its application to clinical medicine’, adding,
30 R.exampDeparBourn31 Da
I have always kept a clinical arm to my activities even though the MRC paid meto do research. The sheep has proved to be an enormously valuable model forwomen—most of the applications which I have made to clinical medicine—ovulationinduction with FSH, pulsatile GnRH, ovarian transplantation and cryopreserva-tion—were all worked out first in the sheep model. (Baird, 2005)
The fragmentation of the SSF/SRF has to some extent been remedied through the orga-nisation of joint meetings with BAS and BFS. For example, the three societies cametogether in April 2005 at the University of Warwick as the ‘UK Fertility Societies’. 300people attended the meeting. The programme included symposia dealing with ‘Geneticsand fertility’, ‘Gamete interactions’, and ‘Influences on Development’; eight other majorlectures (four of which had a clear clinical orientation), a ‘Surgical Symposium’, dealingwith pelvic surgery in people, a session considering the ‘Psychological problems of child-lessness in S.E. Asian communities’, and a nursing section focussed on ‘Human fertility’.There was a very satisfactory balance in the programme between the analysis of clinicalproblems and the experimental investigation of reproductive biology. This was apparentalso in the Short Communications, whether oral or poster presentations. There were118 of these, 40% of which dealt with reproductive problems of livestock, 28% with humanproblems, 24% involved experimental studies of common laboratory animals, and 5% withmore unusual species. 69% of these came from clinical units or laboratories in Britain, 11%from North America, 11% from mainland Europe or Ireland, 5% from Australasia, 3%from Asia and just one from (South) Africa. Such joint meetings are strongly to be encour-aged. In practice reproduction is politically, socially, and physiologically a global problem,requiring research at various levels of biological organisation, from populations (whetherhuman or other animals) through the clinic and whole animals, to the cellular and sub-cellular. The investigation of fertility and reproduction seems to be thriving in Britainat least in this respect: combining the SRF, the BFS and the BAS, produces a membershipof UK Fertility Societies of 1,642. On the other hand the formation in 1985 of the Euro-pean Society for Human Reproduction and Embryology must certainly have divertedclinical and non-clinical scientists from these three societies, which had, by their exam-ple, perhaps contributed to its formation. The total present membership of ESHRE is
G. Edwards, FRS, initially trained in agriculture and later, through various research posts dealing with, forle, mammalian sperm, oocytes and early embryos of experimental animals, became Professor in thetment of Physiology, Cambridge University, and played a very important part in the establishment ofHall, Cambridge, a distinguished centre for Artificial Reproductive Technology. See also n. 21 above.vid Baird, Emeritus Professor of Obstetrics and Gynaecology, University of Edinburgh.
4,500 people, from 101 different countries. Of these 314 are from Britain. Its size is appar-ent from the programme for the 2003 meeting, which runs to more than fifty pages.
Since the creation of SSF/SRF, SFEF, and SRF, the investigation of reproductivephysiology has comprised epidemiology; the study of whole animals and their organs;the measurement in tissues and body fluids of the amounts of hormones, enzymes, andother metabolites; and the investigation by microscopy of events at tissue, cellular orsub-cellular levels, as illustrated by recent work on oocytes, fertilisation, and early stagesof embryonic development. Research carried out at the level of the genome, mitochondria,and ionic exchanges, has led to the modelling of intra-cellular processes, with predictionsthat can be tested experimentally. Aside from the advances in technique to which I havealready alluded, the understanding of a problem central to developmental biology, namelythe roles of different components of the early embryo in its subsequent development, hasbeen greatly increased by the techniques devised by Richard Gardner32 when he was aPh.D. student in R. G. Edwards’s laboratory in Cambridge. This makes possible theremoval from, or addition to, early mammalian embryos of cells or nuclei (Edwards,1980). As well as contributing to fundamental research in embryology, this technique isof fundamental importance in Artificial Reproductive Technology. From its largely unre-marked beginnings (Steptoe & Edwards, 1978; Edwards, 1980) ART is now applied world-wide to problems of fertility in people, as well as to the breeding of livestock, latelyillustrated by the birth of ‘Dolly the Sheep’ (Wilmut & Highfield, 2006). The cloning whichproduced ‘Dolly’, involving such manipulations of oocytes, now attracts much publicinterest, and has even entered into children’s literature (Richardson, 1999).
This paper documents the growth of specialist scientific societies for the study of repro-duction and in doing so provides a framework for understanding the history of this diversefield. I have shown that there were differences between SSF/SRF, the SFEF, and the SSRin their origins. The SSF arose from concern with clinical conditions bearing on fertilityand sterility of people, with some linkage to veterinary and agricultural problems. It thendiversified, embracing reproductive problems in mammals more generally, and encourag-ing a comparative approach. The SFEF at its inception focussed exclusively on human fer-tility, and sustained this for some years before encompassing also livestock and commonlaboratory animals. The SSR emerged seemingly fully formed from a group of scientistsfor the most part interested in the reproductive physiology of livestock, but amongstwhom the aspiration was to consider mammals and other animals more generally. Whatall three societies had in common was that each arose from the initiative, enthusiasmand commitment of a small number of people with particular viewpoints bearing on repro-ductive biology. What is striking about the SSF/SRF is the international character ofthose contributing to its and its journal’s organisation, and the global spread of authorsof papers published in the Journal of Reproduction and Fertility and Reproduction, a fea-ture also of the SSR’s journal, Biology of Reproduction.
Acknowledgements
I am very grateful to Dr Brian Cook (formerly of the Endocrine Unit, UniversityDepartment of Pathological Biochemistry, Royal Infirmary, Glasgow, Scotland),
32 R. L. Gardner, FRS, Edward Penley Abraham Research Professor of the Royal Society, Department ofZoology, Oxford University.
Professor Philip Dziuk (Department of Animal Sciences, University of Illinois, USA), Pro-fessor Patricia Hoyer (Department of Physiology, University of Arizona, USA), and DrKarine Reynaud (Biologie de la Reproduction, Ecole Nationale Veterinaire d’Alfort,France) for their help in the preparation of this contribution. Fig. 1 is reproduced by kindpermission of Christ’s College, Cambridge. Dr Sarah Wilmot and two referees also pro-vided important and useful comments.
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