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Edwards, Sir Robert Geoffrey (Bob)free

(1925–2013)
  • Martin H. Johnson

Sir Robert Geoffrey (Bob) Edwards (1925–2013)

by Anita Corbin and John O'Grady, 1989

© Anita Corbin and John O'Grady; National Portrait Gallery, London

Edwards, Sir Robert Geoffrey (Bob) (1925–2013), physiologist, was born on 27 September 1925 at the maternity home in Bradford Road, Batley, Yorkshire, the son of Samuel Edwards (b. 1896), railway tunnel miner, and his wife Margaret, née Street (b. 1893), a machinist in one of the local mills. He came from a solidly working-class family. His paternal grandfather, Emmanuel Edwards, was a coal miner, born in Portobello, Staffordshire, in 1860. In 1885 he married Eliza Boobyer, two years his junior, and they had sixteen children, all born in Ibstock, Leicestershire, of whom thirteen survived to adulthood, most of the sons becoming miners. Edwards's mother was born in Ardwick, Lancashire. After she married Edwards's father in 1920, they lived in Levenshulme, Manchester, but by the time of Edwards's birth they had moved to 30 Oakhill Road, Batley. Edwards was the middle son, with an older brother, Sammy (b. 1922), and a younger, Harry (b. 1927). When Edwards was five, the family moved to a council house at 25 Highgate Crescent, Gorton, Manchester.

Education, army service, and early career

All three boys were academically able, and were awarded scholarships to Manchester Central Boys' High School. However, Sammy refused to take his up his scholarship, which resulted in heavy maternal pressure on Bob and Harry to do so. Edwards went there in 1937, but his education was interrupted by the Second World War, the school being evacuated to Blackpool. Neither Bob nor Harry liked this and they returned home after six months, and so both were sent by their mother to a farm near Chapel-le-Dale in the Yorkshire Dales to avoid the air raids. Their mother remained in Manchester throughout the war, but the boys saw their father regularly because he was then working on the Settle to Carlisle railway's Blea Moor tunnel. They stayed in Chapel-le-Dale for a year, with no formal education, despite which the two boys learned a great deal about the husbandry of sheep and cattle, and the experience resulted in Edwards's lifelong love of the Yorkshire Dales, to which he would return regularly in later years for family holidays.

On leaving school in 1943, Edwards, who had been known as ‘Geoff’ until he was eighteen, was conscripted for war service in the army; he was trained as a driver-mechanic and served in the Middle East, where he developed his lifelong sympathy for the cause of the Palestinian Arabs. After being demobilised in 1948, he decided to apply to study agricultural sciences at the University College of North Wales, Bangor. By then he was an experienced 23-year-old, described by his impressionable 18-year-old public school-educated and self-described 'unlikely' friend, John Slee, as being 'both ambitious and flexible, and unusually confident in his own judgement' (Slee, 1). However, he found the course disappointingly lacking in intellectual challenge, so for his final year he transferred to zoology, though that year was not enough to salvage his honours degree, and in 1951 he gained a simple pass degree. Temporarily disheartened, he nonetheless applied to study for a postgraduate diploma in animal genetics at Edinburgh University under Conrad Waddington. He clearly impressed Waddington as he offered Edwards, having completed the diploma, a PhD place (1952–5), followed by two years of post-doctoral research, and funded it to the princely sum of £240 per year. It was during this time that Edwards met and married (on 4 September 1956 in Shipton-under-Wychwood parish church, Oxfordshire) his colleague, Ruth Eileen Fowler (1930–2013), endocrinologist, daughter of [Fowler, Sir Ralph Howard], mathematical physicist and weapons researcher, sister of [Fowler, Peter Howard], physicist, and granddaughter of the 1908 Nobel laureate, [Rutherford, Ernest Baron Rutherford of Nelson]. Bob and Ruth Edwards had five daughters between 1959 and 1964: Caroline, Sarah, Jenny, and twins, Anna and Meg.

1955–8 was also the period during which Edwards's scientific flair, curiosity, and industry first became apparent: witness the publication of some thirty-eight papers, many in Nature, each of which concerned, directly or indirectly, genetic aspects of the developmental biology of the mouse. This was a remarkably prescient choice of research subject, given that the human karyotype was only accepted as comprising forty-six chromosomes in 1956, the first demonstration that each cell of the body carried a full set of DNA/genes did not occur until 1958, agreement was reached on the Denver system of classification of human chromosomes in 1960, and the chromosomal aneuploidies underlying developmental anomalies such as Down, Turner, and Klinefelter syndromes were not described until 1959.

Research on oocyte maturation

It was Edwards's interest in trying to understand the origins of the latter syndromes that led him to try to achieve oocyte maturation in vitro, first in mice, then in monkeys, and finally in humans. Thus in a paper published in Nature in 1962 he began: 'Many of the chromosomal anomalies in man and animals arise through non-disjunction or lagging chromosomes during meiosis in the oocyte. Investigation of the origin and primary incidence of such anomalies would be greatly facilitated if meiotic stages etc., were easily available' (Meiosis in ovarian oocytes of adult mammals, Nature, 196 [1962], 446). He started this work in the Experimental Biology Division, headed by Alan Parkes, of the Medical Research Council's National Institute for Medical Research at Mill Hill, London, to which he had relocated in 1958, after a year spent at the California Institute of Technology, primarily to research immunological methods of limiting fertility–then a major scientific goal.

Edwards's work on oocyte maturation, which was initially very much a second string, consisted of simply releasing immature mouse, rat, or hamster eggs from their ovarian follicles into culture medium containing the ovulatory hormone human chorionic gonadotrophin, to explore whether he could simulate their in vivo development. To his amazement he found that the eggs seemed to mature at the same rate as they had in vivo, but did so whether or not the hormone had been added, the eggs evidently maturing spontaneously when released from their follicles. Were the same thing to happen in humans, then the study of the chromosomal dance during human egg maturation was a realistic practical possibility, as was in vitro fertilization (IVF) and thereby studies on the genetics of early human development. However, Edwards's excitement at seeing eggs spontaneously maturing was temporarily blunted by his discovery from reading in the library that Gregory Pincus in the 1930s and Min Chueh Chang in the 1950s had got there before him, using both rabbit and, according to Pincus, human eggs.

Undeterred, Edwards nonetheless pursued his cytogenetic studies on the maturation of human eggs, but was at first delayed by the fact that he required a reliable supply of ovarian tissue from which to retrieve and mature eggs, problematic for a scientist with no medical qualification, given the lack of awareness of this work amongst most of the UK gynaecological profession at that time. However, from 1959 Molly Rose, a gynaecologist at the Edgware General Hospital, near Mill Hill in north-west London, provided biopsied ovarian samples intermittently for the next ten years. At first, Edwards used these biopsies to repeat Pincus's observations from the 1930s, and given the sporadic supply of human material, he also tried monkey and baboon ovarian eggs, but in all cases with little success compared with smaller rodents. In his Nature paper in 1962, Edwards cautiously interpreted the few maturing human (3 of 67), monkey (10 of 56), and baboon (13 of 90) eggs that he had observed as most likely arising from in vivo stimulation and thus partially matured at the time of their recovery from the biopsy. He suggested that Pincus's observation that human eggs require only twelve hours to mature was likely to be artefactual, the source of a confrontation with Pincus some four years later at a conference in Venice.

However, by 1962 Edwards's contract at Mill Hill had come to an end, and so he ceased the work on egg maturation and went for a year to work in the Biochemistry Department at Glasgow University with John Paul, who was then the acknowledged master of tissue culture in the UK and who had learned of some experiments that Edwards had been doing ‘on the side’ at Mill Hill, attempting to generate stem cells from rabbit embryos. The aim of these experiments was to use the stem cells to study early developmental mechanisms, either in vitro, or in vivo after their incorporation into embryos. Together with fellow Glasgow biochemist Robin Cole, they undertook work remarkable in its prescience. Thus they succeeded in producing embryonic stem cells that were capable of proliferating for over 100 generations and of differentiating into a variety of cell types; this work was done some twenty years before Martin Evans and Matthew Kaufman, in 1981, described the derivation of embryonic stem cells from mice. The ignorance of the Glasgow work by those in the stem cell field is probably attributable to the then lack of reliable molecular markers of the various cell types.

Move to Cambridge

At that time the Edwards family was growing, and Ruth had remained in north London with their three young daughters, while Edwards commuted to Glasgow for the working week. Then in 1963, accompanied by his growing family, he arrived in Cambridge as a Ford Foundation Research Fellow at the invitation of Alan Parkes, by then professor of the physiology of reproduction at the university; he was to remain there until his death fifty years later. On arrival, he was funded by the Ford Foundation via grants first to Parkes and then to Bunny Austin (successive heads of the Marshall Laboratory of mammalian reproduction in the Department of Physiology) to continue work on basic reproductive mechanisms, with an eye to developing new methods of fertility control.

In Cambridge Edwards resumed his oocyte maturation experiments, and showed, in a paper published in The Lancet in 1965, that eggs of the human would indeed mature in vitro, but that they simply needed longer than those of smaller mammals, human eggs taking up to thirty-six hours rather than the twelve hours or less erroneously reported by Pincus. However, he had even more difficulty sourcing ovarian biopsy material in Cambridge, and so during the summer of 1965 he turned to the USA for help and so began a long-standing contact with Howard and Georgeanna Jones, then in the Obstetrics and Gynecology Department of the Johns Hopkins University, Baltimore. The supply of American eggs generated during his six-week visit allowed him to confirm the maturation timings that he had published in 1965. However, attempts to fertilize the matured oocytes consistently failed, because of the requirement to ‘capacitate’ the spermatozoa, a final maturation process which spermatozoa undergo physiologically in the uterus, and that is essential for the acquisition of fertilizing competence. Despite exposing sperm to a variety of different treatments aimed at enabling them to acquire fertilizing ability, including, during the summer of 1966 spent at Chapel Hill in North Carolina, USA, an attempt to address this problem by preparing small Millipore chambers loaded with sperm surrounded by a semi-permeable membrane that were then inserted overnight in the uteri of women, thereby permitting prolonged exposure of the sperm to secretions of the female tract, he failed to achieve fertilization convincingly. Progress with in vitro fertilization was thus proceeding very slowly.

Collaboration with Patrick Steptoe and Jean Purdy

By the late 1960s it was clear to Edwards that he needed the services of a UK-based clinician who was sympathetic to his aims and committed to helping him realize them. He found such a person in Patrick Steptoe, an obstetrician who had long been acquainted with the human misery caused by infertility and was also engaged in the pioneering development of relevant laparoscopic (or keyhole) surgical procedures. Edwards initially contacted Steptoe by phone in the autumn of 1967, because he thought the laparoscopic technique might help him recover capacitated human sperm from the oviduct. However, after they met in February 1968, Steptoe, together with other obstetricians, provided sufficient excised ovarian material to Edwards to enable him to embark on a serious attempt to obtain fertilization with oocytes matured in vitro. In this regard, the work of research student Barry Bavister proved decisive; he had found that simply raising the pH of the medium in which the sperm were suspended was sufficient to enable them to acquire fertilizing capacity. Applying the same adjustment to the medium in which human in vitro matured oocytes were mixed with recently ejaculated, washed sperm enabled the early stages of human fertilization to be obtained unequivocally outside the body for the first time. This study was published by Edwards, Steptoe, and Bavister in Nature on 15 February 1969.

Also in 1968, Edwards had recruited Jean Purdy as a technician, one of her attractions being her nursing qualification. Purdy was to assume a key role as the third member of the research team (with Edwards and Steptoe) in the work to come. Together, the three of them formed a committed working team that was to endure from 1968 until Purdy's untimely death aged thirty-nine in 1985 which was followed by Steptoe's just three years later. Before the three met, Edwards's interest in achieving fertilization in vitro was firmly focused on exploring the origin of genetic, particularly chromosomal, disorders of development. Thus, in its discussion, the 1965 Lancet paper, which displayed a breathtaking clarity of vision as Edwards set out a programme of research that predicted the events of the next twenty years and beyond, afforded a heavy focus on the early study and detection of genetic disease compared with the slight emphasis on infertility alleviation. This genetic focus continued in his research papers over the four years to 1969. Thus, in 1968, working with his graduate student Richard Gardner, he published in Nature proof of principle for Preimplantation Genetic Diagnosis (PGD), anticipating the development of PGD clinically by some twenty-two years. Likewise, also in 1968, working with the Cambridge geneticist Alan Henderson, Edwards published (again in Nature) his ‘production line theory’ of egg production to explain the origins of maternal aneuploidy in older women. Thus, the earliest eggs to enter meiosis in the foetal ovary were shown to have more chiasmata and to be ovulated earlier in adult life than those entering meiosis later in foetal life. It was Steptoe, supported by Purdy, who raised the profile for Edwards of IVF as a possible means to alleviate infertility.

The first ‘test tube baby’

The publication of the 1969 paper in Nature immediately placed Edwards and Steptoe in the public limelight in a way that was to set back the future progress of their work. Thus, despite the fact that in the ensuing eighteen months they, together with Purdy, achieved controlled hormonal induction of oocyte maturation in vivo, the laparascopic recovery of these eggs from mature follicles, and early development of the eggs when fertilized to the blastocyst stage in vitro, when the three of them approached the Medical Research Council (MRC) in 1970 to enquire about funding the work, including a move for Steptoe to Cambridge, the referees' reports were very critical of their willingness to talk with the media. One referee began his report: 'Dr Edwards feels the need to publicise his work on radio and television, and in the press, so that he can change public attitudes. I do not feel that an ill-informed general public is capable of evaluating the work and seeing it in its proper perspective. This publicity has antagonised a large number of Dr. Edwards' scientific colleagues, of whom I am one' (National Archives, MRC, papers, NA FD 10/161, referees' reports, 1971).

The grant application was unsuccessful, an unsurprising outcome given the enduring and extraordinarily hostile reactions to the work from many quarters, including strong opposition within the medico-scientific establishment. Thus not only was the MRC against them for the next eight years, the British Medical Association repeatedly portrayed Edwards as Dr Frankenstein, which led him to sue that organization successfully for libel. His early medico-scientific critics included such distinguished figures as Robert Winston, James Watson, Max Perutz, Victor Rothschild, Anne McLaren, and Roger Short, and at times Edwards must have felt very isolated. Indeed, in 1974 he nearly gave up the work, partly because of the lack of progress and the constant opposition, but also because at about that time he had become engaged in politics, and served two terms as a Labour councillor in Cambridge, an experience he enjoyed sufficiently to consider standing as an MP. However, he was persuaded by Purdy and by the desperate plight revealed in the letters from the infertile, to continue. The desperation was real, but in their deliberations it is clear that the MRC did not believe that infertility was a problem worth overcoming, reflecting a common professional viewpoint then, as the letters from patients to Edwards made abundantly clear.

The failure to obtain MRC funding caused a major headache for Edwards and Purdy, who had to collaborate with Steptoe over the considerable distance between Cambridge and Oldham, where Steptoe conducted his clinical work. When a volunteer was available there, one or both of them would have to drive north, often at short notice, for a stay of anything from one day to a week or more; this was difficult for Edwards, given his Cambridge family, research, and teaching responsibilities, and this was where Purdy became so important. This travel, amongst other costs, was supported financially between 1969 and 1978 by an American heiress, Lillian Lincoln Howell, to the tune of at least £89,000.

Edwards, Steptoe, and Purdy were now confronted with the dilemma common to all who wish to translate experimental advances into clinical reality. Given that the embryos obtained in vitro looked normal morphologically, both in the living state and as fixed and stained preparations, was it safe to start placing them in women volunteers with the aim of ‘treating’ their infertility? Such a decision also posed practical problems, such as how to optimize conditions for IVF, embryo culture, and transfer to the genital tract, which day to transfer and by which route, and how to optimize endocrine conditions for inducing ovulation which did not adversely affect implantation. It was the last of these that occupied most of their energy in the long years between the first attempt at transfer in December 1971 and the first successful birth, in July 1978.

Initially, no embryos produced by IVF using gonadotrophin injections to optimize the time and yield of oocyte recovery implanted successfully. Hormonal measurements indicated that these injections had perturbed the luteal phase of the cycle and thereby compromised the development of replaced embryos. When attempts were made to correct these luteal-phase perturbations hormonally, one secure case of further development of an embryo in utero was recorded following a transfer in June 1975, when at eleven weeks after oocyte recovery a tubal pregnancy was diagnosed and terminated. These findings prompted the decision from November 1977 to focus on natural cycles using regular monitoring of urinary oestrogens and luteinizing hormone to detect the surge of the latter hormone that presages ovulation. This, of course, placed a premium on technical efficiency because it meant that only a single mature oocyte was typically available per cycle. It was this approach that proved successful, resulting in the birth of the first 'test tube baby' (as she was described by an enthralled British press), Louise Joy Brown, on 25 July 1978, weighing 5 pounds 12 ounces. Shortly thereafter, in January 1979, a second healthy baby, Alastair MacDonald, was born. These successes were soon followed by IVF births in Australia and the USA, with the procedure rapidly spreading to other countries so that a wealth of experience in improving its efficiency was established over a short time.

This eventual triumph after years of effort, in the face of sustained criticism, did not signal the end of Edwards's problems. The coincidence of the births with Steptoe's retirement from the National Health Service provided an opportunity for Edwards and Purdy to discontinue their arduous travel to and from Oldham by bringing Steptoe to the Cambridge area. However, the university could not offer the required facilities, and public funding for securing them elsewhere was not forthcoming, so their IVF work ceased for more than eighteen months while private sources of support were explored. Eventually, sufficient funding was scraped together to purchase Bourn Hall, a handsome Jacobean house in the Cambridgeshire village of Bourn. Soon a cluster of Portakabins was assembled to provide wards, an operating theatre, and laboratory facilities, work was resumed there in September 1980, and IVF was refined in various ways so that by December 1987 the clinic celebrated the birth of its first 1000 babies. By then Edwards's research had been recognized by his election as a Fellow of the Royal Society in 1984.

Wider professional interests and international recognition

Throughout his extraordinarily productive career, Edwards's interests and energies extended well beyond the confines of the laboratory. Thus, aside from his forays into politics, he was a pioneer in promoting the ethical study of reproduction and public education about science, his catholicity of interests and ideas being matched by his zeal to disseminate them, whether within science, within academe, or amongst the public. His earliest regular publication (from 1969 to 1990) was a news-sheet entitled Research in Reproduction, which he edited. Funded by the International Planned Parenthood Federation and aimed at keeping busy clinicians informed about relevant research, it included reviews of relevant articles and very instructive up-to-date wall charts. Later, Edwards, together with Jean Cohen, a Paris-based obstetrician with whom Edwards had established a close friendship while helping him set up IVF in France, sought support for founding a European counterpart of the American Fertility Society. The two agreed on the then revolutionary idea that science should have equal standing with medicine in such an organization, reflecting science's seminal role in the development of assisted conception and related procedures. This idea was presented by Edwards and Cohen to colleagues at a meeting in Helsinki in 1984, where it was received enthusiastically, and the European Society for Human Reproduction and Embryology (ESHRE) was born in Bonn in 1985 with Edwards as its first chairman, a role to which his boundless enthusiasm and energy were well suited.

The ESHRE, which soon attracted thousands from an impressive number of countries to its annual meetings, also organized numerous workshops covering good laboratory and clinical practice, and produced guidelines on ethical issues. Also discussed at Bonn was the establishment of the Society's own journal, which, under the title Human Reproduction, started publication in 1986 with IRL Press, soon thereafter taken over by Oxford University Press, with Edwards as editor-in-chief. From the outset, he aimed to publish high-quality papers, whilst aiding authors who had difficulty expressing themselves clearly in English, something which took a lot of his time. As the number of high-quality research papers increased, he persuaded the ESHRE to found a second journal for review papers, Human Reproduction Update, of which he was also the principal editor. Finally, as more relevant research at a molecular level was submitted, he successfully made the case for establishing yet a third journal, Molecular Human Reproduction, which he also edited. By the time the five-year contract with Oxford University Press was due for renewal, all three journals had achieved very high standing in the field under his editorship, which, however, ended in 2000 suddenly and acrimoniously; Edwards felt that the ESHRE should publish its three journals in house, but he failed to convince the society's executive committee that such a step was wise, and resigned from the editorship of all three journals.

After his resignation, Edwards used his extraordinarily efficient and very loyal editorial team to found and publish a new journal, Reproductive BioMedicine Online, a bold commitment for a 75-year-old to undertake. With emphasis on topical editorials, rapid publication, and lively debate of contentious issues, and also including lay summaries for the non-specialist reader, this journal also proved an unequivocal success. What was particularly impressive about Edwards's editorship of this journal during its first decade was the extraordinary breadth and quality of his own editorial and other contributions, which ranged through genetics, embryology, ethics, politics, reproductive physiology, endocrinology, sociology, law, stem cells, tissue engineering, circadian rhythms, and cancer. Indeed, that breadth was later reflected by the publication, from 2015, of a sister journal, Reproductive BioMedicine and Society Online.

Edwards was a truly remarkable man: a family man who with his wife Ruth and their five daughters used to make regular visits to the Yorkshire Dales, to which he remained very attached, and where he was delighted to serve as president of the Ribblehead Sheep Show. Indeed, after living during their early years in Cambridge in a modern house on Gough Way, the Edwards family moved to Duck End Farm in the rural village of Dry Drayton, where Edwards could indulge his love of planting deciduous trees and Ruth could tend her herd of Angora goats. As his former colleague Gardner wrote, 'It was a great sadness to his family, and numerous colleagues and friends worldwide, that such an original, energetic, inspiring, and engaging individual should have had to endure a progressively debilitating illness throughout the last five years of his life'. Gardner continued:

It was especially hard that it prevented him from going to Stockholm to receive the 2010 Nobel Prize in Physiology or Medicine in person, which would undoubtedly have proved a particularly gratifying antidote to the enduring hostility he had suffered earlier from people who lacked his vision. However, there were three aspects of the award that made it unusual in present times. First, Edwards was the sole recipient, although it would presumably have been shared with Patrick Steptoe had he still been alive. Second, news of the award was greeted with essentially universal approval within the scientific community. Third, rather than being given for elegant or clever science that might at some time in the future prove beneficial to mankind, it was for tangible clinical progress, namely the birth of several million babies to couples who would otherwise have remained childless. Altogether, his is an extraordinary legacy!

Gardner, 98

Edwards, who received many other honours and awards, was appointed CBE in 1988, and was knighted in 2011, died on 10 April 2013 at the Hilton Park Care Centre in Bottisham, Cambridgeshire, of gastrointestinal haemorrhage, chronic renal disease, and advanced dementia. His funeral was held at Churchill College Chapel and Cambridge crematorium on 26 April 2013, and his ashes were buried at Duck End Farm. His wife Ruth died six months later, and they were survived by their five daughters.

Sources

  • J. Slee, ‘RGE at 25—personal reminiscences’, Reproductive Biomedicine Online, 4/1 (2002), 1
  • M. J. Ashwood-Smith, ‘Robert Edwards at 55’, Reproductive Biomedicine Online, 4/1 (2002), 2–3
  • S. Fishel, ‘RGE at 75’, Reproductive Biomedicine Online, 4/1 (2002), 3–6
  • P. R. Brinsden, ‘Thirty years of IVF: the legacy of Patrick Steptoe and Robert Edwards’, Human Fertility, 12/3 (2009), 137–43
  • M. H. Johnson and others, ‘Why the Medical Research Council refused Robert Edwards and Patrick Steptoe support for research on human conception in 1971’, Human Reproduction, 25/9 (Sept 2010), 2157–74
  • M. H. Johnson, ‘Robert Edwards: Nobel Laureate in physiology or medicine’, Nobel Lecture / Nobel Prize Symposium in Honour of Robert G. Edwards, 7 Dec 2010, www.nobelprize.org/nobel_prizes/medicine/laureates/2010/edwards_lecture.pdf, 6 Sept 2016
  • M. H. Johnson, ‘Robert Edwards: the path to IVF’, Reproductive Biomedicine Online, 23/2 (2011), 245–62
  • R. L. Gardner and M. H. Johnson, ‘Bob Edwards and the first decade of Reproductive Biomedicine Online’, Reproductive Biomedicine Online, 22 (2011), 106–24
  • The Times (11 April 2013)
  • Daily Telegraph (11 April 2013)
  • The Independent (11 April 2013)
  • New York Times (11 April 2013)
  • Nature (16 May 2013)
  • Focus on Reproduction (May 2013), 4–5
  • M. Davies, tribute, Royal College of Gynaecologists website, 1 June 2013, www.rcog.org.uk/globalassets/documents/news/obit_robertedwards.pdf, 6 Sept 2016
  • Human Fertility, 16/2 (2013), 94–6
  • C. L. R. Barratt, ‘MHR celebration issue in tribute to Professor Sir Robert Edwards’, Molecular Human Reproduction, 19/12 (Dec 2013), 783–4
  • J. G. Thompson and R. B. Gilchrist, ‘Pioneering contributions by Robert Edwards to oocyte in vitro maturation’, Molecular Human Reproduction, 19/12 (Dec 2013), 794–8
  • S. G. Hillier, ‘IVF endocrinology: the Edwards era’, Molecular Human Reproduction, 19/12 (Dec 2013), 799–808
  • K. Elder and M. H. Johnson, ‘The Oldham notebooks: an analysis of the development of IVF’, Reproductive Biomedicine and Society Online, 1 (2015), 3–70
  • Memoirs FRS, 61 (2015), 81–102
  • WW (2013)
  • personal knowledge (2017)
  • private information (2017)
  • b. cert.
  • m. cert.
  • d. cert.

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(1849–)