237 robert edwards: nobel laureate in physiology or medicine nobel Lecture/Nobel Prize Symposium in Honour of

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Nobel Lecture/Nobel Prize Symposium in Honour of  

Robert G. Edwards, December 7, 2010




Cambridge University, Cambridge, UK.

Today we are here to celebrate the achievements of Robert Edwards. It is 

however a celebration tinged with sadness. Sadness that Bob himself is not 

well enough to be here in person, so in preparing this lecture, I have tried to 

weave into it some of Bob’s written and spoken words. Sadness also that nei-

ther Patrick Steptoe (1913–1988; Fig. 1) [1] nor Jean Purdy (1946–1985; Fig. 

2) [2], two of his key collaborators, are alive to celebrate with him.


Robert Geoffrey Edwards was born on the 27


 of September 1925 in the 

small Yorkshire mill town of Batley. He arrived into a working-class family, 

the second of three brothers − an older brother, Sammy and a younger, 

Harry. These brothers Bob describes as competitive, “all determined to win 

or, if not to win, to go down fighting” [3]. Bob’s mother, Margaret, was a 

machinist in a local mill. She came originally from Manchester, to where 

the family relocated when Bob was about 5, and where he was educated. In 

those days, bright working class kids could take a scholarship exam at age 

10 or 11 in competition for the few coveted places at a grammar school: the 

potential pathway out of poverty and even to University. All three brothers 

passed the exam, but Sammy decided against Grammar School, preferring 

to leave education as soon as he could to start earning. His mother was  

furious at this wasted opportunity, and so when her two younger sons passed 

the exam, there was no question but that they would continue in education. 

So it was that Bob progressed in 1937 to Manchester Central Boy’s High 

School, which, incidentally, also claims Sir James Chadwick, FRS (1891–

1974), another Cambridge professor and Nobel Laureate (in Physics in 1935 

for discovery of the neutron [4]), as a former pupil. Bob’s summers were 

spent in the Yorkshire Dales, where their mother took her sons to be closer to 

their father’s place of work. There Bob laboured on the farms and developed 

an enduring love for the place. 

*  At the Nobel Prize Symposium in Honour of Robert G. Edwards, Martin H. Johnson delivered this lecture.

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These early experiences were formative for Bob. He became a life–long 

egalitarian, for five years a Labour Party councillor [5], willing to listen to 

and to talk with all and sundry, regardless of class, education, status and 

background. Second, he developed an enduring love for and curiosity about 

natural history and especially the reproductive patterns that he observed 

among the farm’s sheep, pigs and cattle in the Dales. Finally, he took great 

pride in being a ‘Yorkshire man’ – with traditional attributes of affability 

and generosity of spirit combined with no-nonsense blunt-speaking. Indeed, 

following his only meeting with Gregory Pincus (1903–1967) [6] at a confer-

ence in Venice in May 1966, at which Bob, the young pretender, clashed with 

the ‘father of the pill’ over the timing of egg maturation in humans, Bob 

paid Pincus the biggest compliment he could imagine, saying “He would 

have made a fine Yorkshireman!” [7]. 

Figure 2. Jean Purdy (1946–1985) 

(courtesy Barbara Rankin).

Figure 1. Patrick Steptoe (1913–1988) 

(courtesy Andrew Steptoe).

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The aftermath of war was to provide an extended interruption to Bob’s 

education: when he left school in 1943, he was conscripted into the British 

Army for almost four years. To his surprise as someone who was from a 

working class family, he was identified as potential officer material and sent 

on an officer-training course, before being commissioned in 1946 (Fig. 3). 

However, the alien life-style of the officers’ mess was not to his taste and 

served to reinforce his socialist ideals. The years in the army were broken by 

9 months compassionate leave back in the Dales, to which he was released to 

help out when his farmer friend there became ill. So engaged did he become 

in farming life that, after discharge from the army in 1948, he returned 

home to Manchester, from where he applied to read agricultural sciences at 

the University College of North Wales at Bangor.

Figure 3. Bob on National Service,1940s (courtesy Ruth 


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Having gained a place and a grant to fund it, the 6 or so months that 

intervened were occupied in a Government desk job in Salford, Greater 

Manchester, work experience that reinforced the attractions of agricultural 

science. So his disappointment in the course offered at Bangor was acute. By 

that time he was an experienced 23 year old, described by his impressionable 

18 year old public-school educated and self-described “unlikely” friend John 

Slee (Fig. 4), as being “both ambitious and flexible, and unusually confident 

in his own judgement” [8]. And in Bob’s confident judgement, the course 

on offer was not ‘scientific’, and he was bored through two tedious years 

of agricultural descriptions, after which he reported that his teachers were 

“glad to see the back of him” in Zoology for a year. The Zoology Department 

offered a course much more to his style and led by the more intellectually 

challenging Rogers Brambell, FRS (1901–1970) [9]. However, that year was 

not enough to salvage an honours degree, and in 1951, aged 26 he gained 

a simple pass. Unbeknown to him at the time, he was not alone in this 

undistinguished academic embarrassment, as neither “Tibby” Marshall, FRS 

(1878–1949), the founder of the Reproductive Sciences [10], nor Sir Alan 

Parkes, FRS (1900–1990), the first Professor of Reproductive Sciences at 

Cambridge [11], and who was later to recruit Bob there, distinguished them-

selves as undergraduates. In 1951, however, Bob “was disconsolate. It was a 

disaster. My grants were spent and I was in debt. Unlike some of the students 

I had no rich parents… I could not write home, ‘Dear Dad, please send me 

£100 as I did badly in the exams.’” [12]. 

Characteristically, however, Bob’s low spirits did not last long. He learned 

that John Slee had been accepted on a Diploma course in Animal Genetics 

Figure 4. John Slee, 1963 (courtesy Ruth 


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at Edinburgh University under Conrad Waddington, FRS (1905–1975) [13]. 

Bob applied, and, despite his pass degree and to his amazement, he was 

accepted. That summer, he worked in various labouring jobs to earn enough 

to pay his way in Edinburgh [14]. It is tempting to see in these experiences 

of the youthful Bob Edwards consequences for his later approach to life: he 

learned that the accepted hierarchies of organisation and ideas were there to 

be challenged, not simply accepted, and that recovery was possible after what 

might seem the severest of knock-backs. These were lessons that Bob was to 

draw on later in his career. 

In Edinburgh, Bob not only started to map out his scientific career, but 

importantly also met Ruth Fowler (Fig. 5), who was to become his life-long 

scientific collaborator, and whom he was to marry in 1954, their five daugh-

ters following between 1959 and 1964: Caroline, Sarah, Jenny, and twins, 

Anna and Meg. Bob initially found himself somewhat overwhelmed, even 

“intimidated” by Ruth’s august family background. Her father, Sir Ralph 

Fowler, FRS (1889–1944) [15], and her maternal grandfather, Lord Ernest 

Rutherford, FRS (1871–1937) [16], were not only both ‘titled’, but both 

also had the most impressive academic credentials imaginable. Ralph Fowler 

was Plummer Professor of Mathematical Physics in Cambridge from 1932 to 

1944, whilst Rutherford was the first Nobel Laureate in Ruth’s family, having 

been awarded the 1908 Nobel Prize in Chemistry ‘for his investigations 

into the disintegration of the elements, and the chemistry of radioactive 


Figure 5. Ruth Fowler in laboratory, Edinburgh, 1950s (courtesy Ruth Edwards).

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The intellectual spirit of scientific enquiry that Bob experienced in 

Edinburgh obviously fitted his aptitudes perfectly, for Waddington rewarded 

his Diploma year with a three year PhD place and funded it with the princely 

sum of £240.00 per year [17]. Bob’s chosen field of research was the develop-

mental biology of the mouse. Bob saw that to understand development 

involved engaging in an interdisciplinary mix, not just of embryology and 

reproduction, the conventional view at the time, but also of genetics. Given 

the scientific and social emphasis on genetics over the last 40 or so years, it 

is difficult now to realise how advanced a view this was in the 1950s, when 

genetic knowledge was still rudimentary and largely alien to the established 

developmental and reproductive biologists of the day, as Bob himself was 

later to comment [18]. For example, it was in the 1950s that DNA was estab-

lished as the molecular carrier of genetic information [19–22], that it was 

first demonstrated that each cell of the body carried a full set of DNA/genes 

[23–25], and that genes were selectively expressed as mRNA to generate  

different cell phenotypes [26]. Perhaps of greater importance for Bob at that 

time, it was only by the late 1950s that cytogenetic studies led to the accepted 

human karyotype as 46 chromosomes [27–28], that agreement was reached 

on the Denver system of classification of human chromosomes [29], and that 

the chromosomal aneuploidies underlying developmental anomalies such as 

Down, Turner and Klinefelter Syndromes were described [30–32]. 

Figure 6. Alan Gates at a meeting in Cambridge in the late 

1950s (possibly 1957).

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Bob worked under his supervisor, Alan Beatty, to generate haploid, 


triploid and aneuploid mouse embryos and studied their potential for normal 

development. In order to undertake what were, in effect, early attempts at 

‘genetic engineering’ in mammals, he needed to be able to manipulate the 

chromosomal composition of eggs, sperm and embryos. Whilst in mice, 

sperm were abundant, eggs were not, and overcoming this deficit led him 

to two major discoveries that proved to be of later significance. First, with 

Ruth, they worked to devise ways of increasing the numbers of synchronised 

eggs recoverable from adult female mice through a series of papers on the 

control of ovulation induced by use of exogenous hormones [33]. In doing 

so, they overturned the conventional wisdom that super-ovulation of adult 

females was not possible. Second, working with an American post-doc, Alan 

Gates (Fig. 6), Bob described the remarkable timed sequence of egg chromo-

somal maturation events that led up to ovulation after injection of the ovula-

tory hormone (human chorionic gonadotrophin; hCG) [34]. His six years,  

between 1951 and 1957, in Edinburgh give an early taste of his prodigious 

energy, resulting in 38 papers. Indeed so productive was this period that the 

last of the papers resulting from his Edinburgh work did not appear in print 

until 1963.

It was also in Edinburgh that Bob’s interest in ethics was first sparked 

by the interdisciplinary debates among scientists and theologians that 

Waddington organised, and, as a result, Bob went on what he describes as a 

“church crawl”, trying the ten of so variants of Christianity on offer in 1950s 

Edinburgh. He did not emerge from his consumer testing “God-intoxicated” 

[35], but convinced that man held his own future in his own hands. Bob’s 

humanist ethical sympathies were to be developed further in all his later 



These early 1950s studies in science and ethics were to form the platform 

on which Bob’s later IVF work was to be based, but before that his interests 

and life took a diversion to the California Institute of Technology for the 

year 1957–8. Bob describes his year at CalTech as being “a bit of a holiday”, 

but it was a holiday which, with hindsight, had distracting consequences. He 

went there to work with Albert Tyler (1906–1968) [36], an influential elder 

statesman of American reproductive science, working on sperm–egg interac-

tions. CalTech was then a hotbed of developmental biology, and Tyler had 

clustered around him an exciting group of young scientists, which included 

that year a visit by the English doyen of fertilisation, Lord Victor Rothschild, 

FRS (1910–1990) [37], who was later to clash scientifically with Edwards over 

his IVF work [38]. In this clash, needless to say, the younger man triumphed 

[39], just as he had with Pincus. Tyler was exploring the molecular specific-

ity of egg–sperm interactions and had turned as a model to immunology. 

Immunology was then at a very exciting phase in its development, with the 

engaging Sir Peter Medawar, FRS (1915–1987, Nobel Laureate in Physiology 

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or Medicine, 1960) [40], influentially for Bob, extending his ideas on im-

munological tolerance to the paradox of the ‘fetus as an allograft’: a semi-

paternal graft nonetheless somehow protected from maternal immune 

attack inside the mother’s uterus. This confluence of reproduction and 

immunology excited Bob’s restless curiosity and hence the choice of Tyler. 

The subject also offered funding possibilities via the Ford and Rockefeller 

Foundations and the Population Council, which were increasingly 


concerned about world population growth and the need for better methods 

to control fertility. Immuno-contraception then seemed to offer tantalisingly 

specific possibilities.

So when Bob returned to the UK from CalTech in 1958 at Alan Parkes’ 

invitation to join him at the Medical Research Council (MRC) National 

Institute for Medical Research (NIMR) at Mill Hill in north London, it was to 

work on the science of immuno-contraception [5]. This period in the USA 

initiated a series of 24 papers on the immunology of reproduction between 

1960 and 1976. It also prompted Bob’s first involvement in founding an 

international society in 1967 in Varna, Bulgaria when the International 

Coordinating Committee for the Immunology of Reproduction was created 

[41]. It was, in retrospect, to prove a distracting diversion from what was 

to become Bob’s main work, albeit one that continued to enthuse Bob for 

many years, witnessed not least by my own recruitment to enter this field 

of study with him as a graduate student in 1966. Nonetheless, the period at 

Mill Hill, between 1958 and 1962, seems to have been a period of increasing 

intellectual conflict for Bob. Whilst enthusiastically working on the science 

underlying immuno-contraception, his old interests in eggs, fertilisation 

and, in particular, the genetics of development were gradually reasserting 

themselves. His day job was therefore increasingly supplemented by evening 

and weekend flirtations with egg maturation.


Bob claims that the stimulus reawakening his interests in eggs was provided 

by the then recent consensus about the number of human chromosomes 

and, more particularly the descriptions in 1959 of the pathologies in man 

that resulted from chromosomal anomalies [42]. Might these anomalies 

result from errors in the complex chromosomal dance that he and Alan 

Gates had observed in maturing mouse eggs? The possible clinical relevance 

of his work on egg maturation and aneuploidy in the mouse was becoming 


So Bob resumed his experimenting with mice, trying to mimic in vitro the 

in vivo maturation of eggs. He tried releasing the immature eggs from their 

ovarian follicles into culture medium containing the ovulatory hormone 

hCG, to see whether he could simulate their in vivo development. Amazingly 

he found it worked the first time: but it did so whether or not the hormone 

had been added. It seemed that the eggs were maturing spontaneously when 

released from their follicles. And the same happened in rats and hamsters. 

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If this also happened in humans, then the study of the chromosomal dance 

during human egg maturation was a realistic practical possibility, as was in 

vitro fertilisation and thereby studies on the genetics of early human develop-

ment. However, Bob’s excitement at seeing eggs spontaneously maturing 

was temporarily blunted by his discovery that Gregory Pincus in the 1930s 

[43–44] and M.C. Chang (1908–1991) [45–46] in the 1950s had been there 

before him, using both rabbit and, Pincus claimed, human eggs. 

In order to pursue his basic science studies on maturation, he needed a 

reliable supply of human ovarian eggs. This requirement posed difficulties 

for a scientist with no medical qualifications, given the elitist attitudes and 

scientific illiteracy then prevalent amongst most of the UK’s gynaecologists. 

His break-through came initially with Molly Rose, at whose door in the 

nearby Edgeware General Hospital he arrived after a recommendation from 

a fellow kindred spirit in John Humphrey, FRS (1915–1997) [47], ten years 

Bob’s senior and the medically qualified Head of Immunology at Mill Hill. 

Notwithstanding his more privileged social background, Humphrey shared 

Bob’s passion for science, its social application and utility, and his left wing 

politics – indeed he had been a Marxist until 1940. Bob asked John if he 

knew anyone who might be helpful, and John suggested Molly Rose and  

offered to arrange an introduction. So off Bob went, and Molly Rose 


provided human ovarian biopsy samples intermittently for the next ten years.

Between 1960 and 1962, Bob tried to repeat and extend Pincus’ observa-

tions, using not only human but also dog, monkey and baboon eggs, but with 

such limited success compared with smaller rodents that in a 1962 Nature 

paper [48], he carefully interprets the few maturing human and baboon eggs 

that he observed as artefacts. But by this time, Bob’s quest for human eggs, 

and his dreams of IVF and studying early aneuploidies in human embryos, 

had reached hostile ears, most notably those of the then Director of the 

Institute, Sir Charles Harington, FRS (1897–1972), who banned any work 

on human IVF at NIMR [49]. Alan Parkes was no longer able to defend Bob, 

having left in 1961 to take up his chair in Cambridge and, although he had 

asked Bob to join him, there was no post until 1963. By the time Bob left 

Mill Hill in 1962 for a year in Glasgow, he had encountered just a taste of the 

opposition to come.


Bob had been invited to Glasgow University’s Biochemistry Department by 

John Paul, then the acknowledged master of tissue culture in the UK, who 

had heard of Bob’s attempts to generate stem cells from rabbit embryos [18]. 

The invitation was to result in a paper [50] remarkable for its prescience – 

the first of eight landmark papers (Table 1) that I identify in this contribution. 

It describes the production of embryonic stem cells from rabbit embryos – 

capable of proliferating through over 100 generations and of differentiating 

into various cell types. This report was published some 18 years before Evans 

and Kaufman described the derivation of ES cells from mouse embryos [51]. 

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That this work has largely been ignored by those in the stem cell field is prob-

ably mainly attributable to its being too far ahead of its time. Thus, reliable 

molecular markers for different types of cells were not available then, nor 

were appropriate techniques with which to critically test the developmental 

potential of the cultured cells.

Table 1. The eight landmark papers

1. Cole 


Edwards R.G., Paul J. (1965) Cytodifferentiation in cell colonies 

and cell strains derived from cleaving ova and blastocysts of the rabbit. 

Exp. Cell Res. 37: 501–4.

2. Edwards 


(1965) Maturation in vitro of human ovarian oocytes. 

Lancet 286: 926–9.

3. Gardner 


Edwards R.G. (1968) Control of the sex ratio at full 

term in the rabbit by transferring sexed blastocysts. Nature 218: 


4. Edwards 


Bavister B.D., Steptoe P.C. (1969) Early stages of fertili-

zation in vitro of human oocytes matured in vitro. Nature 221: 632–5.

5. Steptoe 


Edwards R.G. (1970) Laparoscopic recovery of preovu-

latory human oocytes after priming of ovaries with gonadotrophins. 

Lancet 295: 683–9.

6. Steptoe 


Edwards R.G., Purdy JM. (1971) Human blastocysts 

grown in culture. Nature 229: 132–3.

7. Edwards 


Sharpe DJ. (1971) Social values and research in human  

embryology. Nature 231: 87–91.

8. Steptoe 


Edwards R.G. (1978) Birth after the reimplantation of a  

human embryo. Lancet 312: 366.

Bob arrived in Cambridge from Glasgow in 1963. He describes how he 

immediately reacted against the then extant “misogynist public-school tradi-

tions; the exclusivity…; the privileges given to the already privileged”. But he 

set against that the “sheer beauty of the place… the concern with the truth 

and high seriousness… the ambience of scientific excellence… I was sur-

rounded by so many talented young men and women.” [52] He continued to 

pursue both the immunology of reproduction and egg maturation, working 

furiously on the latter to collect pig, cow, sheep, the odd monkey and some 

human eggs. Eventually, he was able to show that eggs of all these species 

would indeed mature in vitro, but that the eggs of larger animals simply 

needed longer than those of smaller ones, human eggs taking some 36 hours 

rather than the 12 or less hours erroneously reported by Pincus [44]. These 

cytogenetic studies were reported in two seminal papers in 1965 [53–54], 

both of which are primarily concerned with understanding the kinetics of 

the meiotic chromosomal events. As the second landmark paper, I have selected 

the one in The Lancet, in which Bob’s breath-taking clarity of vision is evident 

as he sets out a programme of research that predicted the events of the next 

20 years and beyond (Table 2). You will notice the heavy focus on the early 

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study and detection of genetic disease compared with the slight emphasis on 

infertility alleviation, unsurprising given Bob’s research interests. Indeed, 

within three years he had, with my fellow graduate student Richard Gardner, 

provided proof of principle for preimplantation genetic diagnosis (PGD), 

in a paper on rabbit embryo sexing published in 1968 [55] and my third 

landmark paper. Later, in the 1980s, Bob was to play a key role in promoting 

the development of PGD clinically [56], and PGD was to prove a powerful 

political tool in convincing the UK Parliament to permit research on human 

embryos. And then by 1969 he had reported the first step towards PGD in 

humans by describing IVF [57] – a fourth landmark paper in as many years.

Table 2. Key points in the programme of research laid out in the Discussion to 

Edwards’ 1965 Lancet paper (landmark paper 2) [54].

1.  Studies on non-disjunction of meiotic chromosomes as a cause of 

aneuploidy in humans.

2.  Studies on the effect of maternal age on non-disjunction in relation 

to the origins of trisomy 21.

3.  Use of human eggs in IVF.

4.  Culture of fertilised human eggs in vitro.

5.  Use of priming hormones to increase the number of eggs per wom-

an available for study/use.


Study of early IVF embryos for evidence of (ab)normality – especially 

aneuploidies arising prior to or at fertilisation.

7.  Control of some of the genetic diseases in man.

8.  Control of sex-linked disorders by sex detection at blastocyst stage 

and transfer of only female embryos.

9.  Para-cervical transfer of IVF embryos into the uterus.

10.  Use of IVF embryos to circumvent blocked tubes.

11.  Avoidance of a multiple pregnancy (as observed after hormonal 

priming and in vivo insemination) by transfer of a single IVF embryo.


Underlying both the 1965 and 1969 papers are two scientific struggles: the 

first being simply but critically the continuing difficulty in obtaining a regular 

supply of ovarian tissue. Local Cambridge sources proved unreliable, and 

Molly Rose was now 2–3 hours’ drive away in north-west London, so during 

the summer of 1965, Bob turned to the USA for help and initiated his now 

famous contacts with Howard and Georgeanna Jones [58], then at the Johns 

Hopkins Medical School in Baltimore. This supply of American eggs allowed 

Bob to confirm the maturation timings published in 1965 [54]. However, it 

was the second struggle that was by then occupying most of his attention, 

namely that in order to fertilise these in vitro matured eggs, he had to 


‘capacitate’ the sperm. ‘Capacitation’ is a final maturation process, which 

sperm will usually undergo physiologically in the uterus, and that is essential for 

the acquisition of fertilising competence. Failing to achieve this convincingly 

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at Johns Hopkins, he made a second transatlantic summer journey in 1966 to 

visit Luther Talbot and his colleagues at Chapel Hill. Bob applied his usual 

ingenuity to try a variety of ways to overcome the problem of sperm capacita-

tion, but no reliable evidence for success was forthcoming [59, 60]. Then in 

1968 both struggles began to resolve.

Resolving the problem of sperm capacitation was the initial attraction to 

Bob of Patrick Steptoe’s laparoscopic technique, Bob seeing it as a way of  

recovering capacitated sperm from the oviduct [61]. However, the actual 

solution to this problem lay nearer home. Parkes had retired as Professor in 

1967, to be replaced by Colin ‘Bunny’ Austin (1914–2004) [62] (Fig. 7). In the 

early 1950s, Bunny, and independently M.C. Chang [45], had discovered the 

requirement for sperm capacitation [63–64], and so Bunny set his graduate 

student, Barry Bavister (1943–), to work to try and resolve how to reliably 

capacitate hamster sperm in vitro. Bavister demonstrated a key role for pH in 

a short paper published in 1969 that showed how higher rates of fertilisation 

could be obtained by simply increasing the alkalinity of the medium [65]. Bob 

seized on this observation and co-opted Barry to his project of capacitating 

human sperm. That proved to do the trick, leading to the 1969 paper [57].

The problem of the intermittent egg supply in the UK was also resolving. 

Bob continued to rely on surgeons to provide him with ovarian biopsies 

from which to mature eggs in vitro, indeed four are thanked in the 1969 

Nature paper. According to Bob [66], Molly Rose provided the first group 

of eggs to be fertilised, and although invited to be a co-author, declined for 

reasons unknown. Also thanked are Norman Morris [67], Janet Bottomley 

and Sanford Markham, and although it is not known whether they provided 

any of the ovarian eggs for in vitro maturation described in the paper, Patrick 

clearly did so [66], and at last provided for Bob a potentially more stable 

clinical partnership.

The 1969 Nature paper describing IVF in humans [57] makes modest 

claims, only two of 56 eggs reaching the two-pronuclear stage. But, like Bob’s 

Figure 7. Bob with Bunny Austin, 1960s 

(courtesy Ruth Edwards).

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other papers, it is a model of clarity, describing well-controlled experiments, 

cautiously interpreted. This paper convinced where previous claims [68–73] 

had failed, precisely because the skilled hands and creative intellect that lay 

behind it are so evident from its text. With its publication, announced to the 

media on St Valentine’s Day [74], all hell was let loose. 


So 1969 seemed to Bob to be a good year. Not only did IVF succeed at long 

last, and his partnership with Patrick seemed set to flourish, but also so  

impressed were the Ford Foundation with Bob’s work that they paid for him 

to be awarded a Ford Foundation Readership (a half way step to a professor-

ship) in the University. Elated by his promotion and their achievement, Bob 

and Patrick pressed on, the latter’s laparoscopic skills coming to the fore, 

first in 1970 with the collection of in vivo matured eggs from follicles after 

mild hormonal stimulation [75], and then achieving regular fertilisation of 

these eggs and their early development through cleavage to the blastocyst 

stage [76] – my fifth and sixth landmark papers. So well was the work going 

that in February 1971 they confidently applied to the UK Medical Research 

Council for funding to bring Patrick to Cambridge from Oldham General 

Hospital in Greater Manchester, where Patrick worked [77].

However, any illusions that Bob may have had that their achievements 

would prove a turning point in his fortunes were soon shattered, and just 2 

months later on April Fool’s Day 1971 the MRC decided to reject the grant 

application [77]. The practical consequences of this rejection were profound 

– both psychologically and physically – not least that for the next 7 years, Bob 

shuttled on the 12 hour round trip between Cambridge and Oldham, leaving 

Ruth and his five daughters in Cambridge.

The professional attacks on Bob and his work took a number of forms 

[77], and one must try to make a mental time trip back to the 1960s/70s 

to understand their basis. Despite the nature of the political and religious  

battles to come, his scientific and medical colleagues did not focus on the 

special status of the human embryo as an ethical issue. However, ethical 

issues were raised professionally, but took quite a different form. Thus, it 

is difficult now to comprehend the complete absence of infertility from 

the consciousness of most gynaecologists in the UK at the time, to which 

Patrick Steptoe was a remarkable exception [78]. Indeed, Bob’s strong 

commitment to treating infertility came to the fore only after he teamed up 

with Patrick, his previous priority being the study and prevention of genetic 

and chromosomal disorders. In the several reports from the Royal College 

of Obstetricians and Gynaecologists and the MRC during the 1960s examin-

ing the areas of gynaecological ignorance that needed academic attention,  

infertility simply did not feature [77]. Overpopulation and family planning 

were seen as dominant concerns and the infertile were ignored as at best a 

tiny and irrelevant minority and at worst as a positive contribution to popula-

tion control. This was a values system that Bob simply could not accept, 

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and the many encouraging letters Bob was to receive from infertile couples 

provided a major stimulus to his continued work later, despite so much 

professional and press antagonism and so many set-backs. For his medico-

scientific colleagues, however, the fact that infertility was not seen as a clinical 

issue, meant that any research designed to alleviate it was not viewed as 

experimental treatment, but as using humans for experiments. Given the 

sensitivity to Nazi ‘medical experiments’, and the public reaction and 

disquiet surrounding the recent publication of ‘The Human Guinea-pig’ [79], 

this distinction was critical. The MRC, in rejecting the grant application, took 

the position that what was being proposed was human experimentation, and 

so were very cautious, emphasising risks rather than benefits, of which they 

saw few if any [77].

Bob and Patrick were also attacked for their willingness to talk with the 

media. It is even more difficult nowadays, when the public communication of 

science is so embedded institutionally, to understand how damaging to them 

this was. The massive press interest of the late 1960s was unabated in the 

ensuing years, and so Bob was faced with a choice: either he could keep his 

head down and allow press fantasies and speculations to go unanswered and 

unchallenged, or he could engage, educate and debate. For Bob this was no 

choice, regardless of the consequences for him professionally. His egalitarian 

spirit demanded that he trust to common people’s common sense. His radical 

political views demanded that he fought the corner of the infertile, the 

underdog with no voice. The Yorkshireman in him relished engagement in 

the debate and argument. In the seventh landmark paper selected, published 

in Nature in 1971 with Dave Sharpe [80], he acknowledges the risk to his own 

interests of so doing. Risky it clearly was, one of the scientific referees on 

their MRC grant application starting his referee’s report declaring his strong 

distaste for all the media exposure [77]. Bob was a pioneer in the public 

communication of science, and paid a heavy price for being so.

The Edwards and Sharpe paper [80] is a tour de force in its survey of the  

scientific benefits and risks of the science of IVF, in the legal and ethical 

issues raised by IVF, and in the pros and cons of the various regulatory 

responses to them. It sets out the issues succinctly and anticipates social  

responses that were some 13–19 years into the future. In subsequent years, 

Bob built on his strong commitment to social justice based on a social ethic, 

as he engaged at every opportunity with ethicists, lawyers and theologians,  

arguing, playing ‘devil’s advocate’ (literally, in the eyes of some), and engaging 

in what we would now call practical ethics, as he hammered out his position 

and felt able to fully justify his instincts intellectually.

Edwards_lect_.indd   16

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But all this was to little avail. Indeed, Bob was continually frustrated at the 

unwillingness of most of the establishment to engage seriously in ethical  

debates in advance of the final validation of IVF that was to come in 1978 

with the birth of Louise Brown (Fig. 8), and my eighth and final landmark 

paper [81]. Only then did most UK social hierarchies, such as the MRC, the 

British Medical Association, the Royal Society and Government move gradu-

ally from their almost visceral reactions against IVF and its possibilities to 

serious engagement with the issues [56]. Then, to their credit, both the MRC 

and the Thatcher Government of the time came on board, but it was not 

until 1989, 24 years after Bob’s 1965 landmark paper in the Lancet, that the 

UK Parliament finally gave its stamp of approval to his visionary work, and 

then only after a fierce battle lasting some 11 years [82]. Eleven years since 

the eighth and final landmark paper was published announcing the birth of 

Louise Brown. And of course, it has taken 45 years since that 1965 paper for 

us to be celebrating the award of the Nobel Prize to this remarkable man.

Figure 8. Louise Brown holding the 

1000th Bourn Hall baby, 1987 (courtesy 

Bourn Hall Clinic).

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The eight landmark papers I have selected (Table 1) present us with a 

man of vision and foresight, imagination and intellectual rigour, and extra-

ordinary energy and drive: witness his prodigious output of papers between 

1954–2008 [83]. A man who could inspire colleagues to tread with him on 

a difficult scientific path besieged by public and professional animosity, of 

whom two in particular, Patrick and Jean, we miss today (Fig. 9). And these 

papers do not even begin to touch Bob’s other academic and personal quali-

ties, so evident in the way he has founded and generously nurtured journals 

and international societies [84, 85], transforming the intellectual landscape 

not just of gynaecology, but also of ethics and social anthropology. Truly the 

‘father of Assisted Reproductive Technology’ in its widest interdisciplinary 

sense: the only sense in which Bob knows it. For Bob, it is truly the Nobel 

Prize for Physiology AND Medicine: there can be no OR about it.


I thank the Edwards family for their help in writing this account, for which 

however I take full responsibility. I also thank Kay Elder and Sarah Franklin 

for their unfailing wisdom and helpful advice. I thank Andrew Steptoe for 

permission to reproduce Figure 1, Barbara Rankin for permission to repro-

duce Figure 2, Ruth Edwards for permission to reproduce Figures 3–7, and 

Bourn Hall Clinic for permission to reproduce Figures 8 and 9. The research 

underpinning this account was supported by a grant from The Wellcome 

Trust [088708], which otherwise had no involvement in the research or its 

publication. This chapter is adapted from “Robert Edwards: the path to 

IVF” by M.H. Johnson published in Reproductive BioMedicine Online (Elsevier)  

at doi:10.1016/j.rbmo.2011.04.010 and is included in this volume with 



Figure 9. Bob, Jean and Patrick at Bourn 

Hall, 1981 (courtesy Bourn Hall Clinic).

Edwards_lect_.indd   18

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  1.  Edwards, R.G., “Patrick Christopher Steptoe, C. B. E. 9 June 1913–22 March 1988,” 

Biog Mems Fell R Soc, 1996. 42: p. 435–452.

  2.  Edwards, R.G. and P.C. Steptoe, “Preface,” In: Edwards, R.G., Purdy, J.M., Steptoe, 

P.C. (eds). Implantation of the Human Embryo, 1985. Academic Press: London, UK,  

p. vii–viii.

  3.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 25.

  4.  Massey, H. and N. Feather “James Chadwick. 20 October 1891 – 24 July 1974,” Biog 

Mems Fell R Soc, 1976. 22: p. 10–70.

 5.  Ashwood-Smith, M.J., “Robert Edwards at 55,” Reprod BioMed Online, 2002. 4 

(Suppl.1): p. 2–3.

  6.  Ingle, D.J., “Gregory Goodwin Pincus. April 9,1903–August 22,196,” Biog Mems Natl 

Acad Sci, 1971: p. 229–270.

  7.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 43.

 8.  Slee, J., “RGE at 25 – personal reminiscences,” Reprod BioMed Online, 2002. 4 

(Suppl.1): p. 1.

  9.  Oakley, C.L., “Francis William Rogers Brambell. 1901–1970,” Biog Mems Fell R Soc

1973. 19: p. 129–171.

10.  Parkes, A.S., “Francis Hugh Adam Marshall. 1878–1949,” Biog Mems Fell R Soc, 1950. 7: 

p. 238–251.

11.  Polge, C., “Sir Alan Sterling Parkes. 10 September 1900 – 17 July 1990,” Biog Mems Fell 

R Soc, 2006. 52: p. 263–283.

12.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 7.

13.  Robertson, A., “Conrad Hal Waddington. 8 November 1905 – 26 September 1975,” 

Biog Mems Fell R Soc, 1977. 23: p. 575–622.

14.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 18.

15.  Milne, E.A., “Ralph Howard Fowler. 1889–1944,” Biog Mems Fell R Soc, 1945. 5: 


p. 60–78.

16.  Eve, A.S. and Chadwick, J., “Lord Rutherford. 1871–1937,” Biog Mems Fell R Soc, 1938. 

2: p. 394–423.

17.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 20.

18.  Edwards, R.G., “An astonishing journey into reproductive genetics since the 1950’s,” 

Reprod Nutr Dev, 2005. 45: p. 299–306.

19.  Watson, J.D. and F.H. Crick, “Genetical implications of the structure of deoxyri-

bonucleic acid,” Nature, 1953. 171: p. 964–967.

20.  Watson, J.D. and F.H. Crick, “Molecular structure of nucleic acids: a structure for 

deoxyribose nucleic acid,” Nature, 1953. 171: p. 737–738.

21.  Franklin, R. and R. Gosling, “Molecular configuration in sodium thymonucleate,” 

Nature, 1953. 171: p. 740–741.

22.  Wilkins, M.H.F., A.R. Stokes and H.R. Wilson, “Molecular structure of deoxypentose 

nucleic acids,” Nature, 1953.171: p. 738–740.

23.  Gurdon, J.B., “Adult frogs derived from the nuclei of single somatic cells,” Dev Biol, 

1962. 4: p. 256–273.

24.  Gurdon, J.B., “The developmental capacity of nuclei taken from intestinal epithelium 

cells of feeding tadpoles,” Development, 1962. 10: p. 622–640.


25.  Gurdon, J.B., T.R. Elsdale and M. Fischberg, “Sexually mature individuals of Xenopus 

laevis from the transplantation of single somatic nuclei,” Nature, 1958. 182, p. 64–65.

26.  Weinberg, A.M., 2001. “Messenger RNA: origins of a discovery,” Nature, 2001. 414:  

p. 485.

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27.  Tjio, J.H. and A. Levan, “The chromosome number of man,” Hereditas, 1956. 42: 


28.  Ford, C.E. and J.L. Hamerton, “The chromosomes of man,” Nature, 1956. 178: 


p. 1020–1023.

29.  Denver Conference, “A proposed standard system of nomenclature of human mitotic 

chromosomes,” Lancet, 1960. 275: p. 1063–1065.

30.  Ford, C.E., P.E. Polani, J.H. Briggs and P.M. Bishop, “A presumptive human XXY/XX 

mosaic,” Nature, 1959. 183: p. 1030–1032. 

31.  Ford, C.E., K.W. Jones, P.E. Polani, J.C. De Almeida and J.H. Briggs, “A sex-chromo-

some anomaly in a case of gonadal dysgenesis (Turner’s syndrome),” Lancet, 1959. 

273: p. 711–713. 

32.  Lejeune, J., M. Gautier and R. Turpin, “Etude des chromosomes somatiques de neuf 

enfants mongoliens,” Comptes Rendus Hebd Seances Acad Sci, 1959. 248: p. 1721–1722. 

33.  Fowler, R.E. and R.G. Edwards, “Induction of superovulation and pregnancy in 


mature mice by gonadotrophins,” J Endocr, 1957. 15: p. 374–384. 

34.  Edwards, R.G. and A.H. Gates, “Timing of the stages of the maturation divisions, 

ovulation, fertilization and the first cleavage of eggs of adult mice treated with 


gonadotrophins,” J Endocr, 1959. 18: p. 292–304.

35.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 23–4.

36.  Horowitz, N.H., C.B. Metz, J. Piatigorsky, L. Piko, J.D. Spikes, and M. Ycas, “Albert 

Tyler,” Science, 1969. 163: p. 424.

37.  Reeve, S., “Nathaniel Mayer Victor Rothschild, G.B.E., G.M. Third Baron Rothschild. 

31 October 1910 – 20 March 1990,” Biog Mems Fell R Soc, 1994. 39: p. 364–380.

38.  Rothschild, “Did fertilization occur?’ Nature, 1969. 221: p. 981.

39.  Edwards, R.G., B.D. Bavister and P.C. Steptoe, “Did fertilization occur?” Nature, 1969. 

221: p. 981–982.

40.  Mitchison, N.A., “Peter Brian Medawar. February 1915 – 2 October 1987,” Biog Mems 

Fell R Soc, 1990. 35: p. 282–301.

41.  Rukavina, D., “The history of reproductive immunology: my personal view,” Am J 

Reprod Immunol, 2008. 59: p. 446–450.

42.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 38.

43.  Pincus, G. and E.V. Enzmann, “The comparative behavior of mammalian eggs in vivo 

and in vitro i. the activation of ovarian eggs,” J Exp Med, 1935. 62: p. 665–675.

44.  Pincus, G. and B. Saunders, “The comparative behavior of mammalian eggs in vivo 

and  in vitro. VI. The maturation of human ovarian ova,” Anat Record, 1939. 75: p. 


45.  Greep, R.O., “Min Chueh Chang. October 10, 1908 — June 5, 1991,” Biog Mems Natl  

Acad Sci. http://www.nap.edu/readingroom.php?book=biomems&page=mchang.html

46.  Chang, M.C., “The maturation of rabbit oocytes in culture and their maturation,  

activation, fertilization and subsequent development in the Fallopian tubes,” J Exp 

Zool, 1955. 128: p. 379–405.

47.  Askonas, B.A., “John Herbert Humphrey. 16 December 1915–25 December 1987,” 

Biog Mems Fell R Soc, 1990. 36: p. 274–300.

48.  Edwards, R.G., “Meiosis in ovarian oocytes of adult mammals” Nature,  1962. 196:  

p. 446–450.

49.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 48.

50.  Cole, R.J., R.G. Edwards and J. Paul, “Cytodifferentiation in cell colonies and cell 

strains derived from cleaving ova and blastocysts of the rabbit,” Exp Cell Res, 1965. 37: 

p. 501–504. 

51.  Evans, M.J. and M.H. Kaufman, “Establishment in culture of pluripotential cells from 

mouse embryos,” Nature, 1981. 292: p. 154–156.

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52.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 51.

53.  Edwards, R.G., “Maturation in vitro of mouse, sheep, cow, pig, rhesus monkey and  

human ovarian oocytes,” Nature, 1965. 208: p. 349–351.

54.  Edwards, R.G., “Maturation in vitro of human ovarian oocytes,” Lancet, 1965. 286:  

p. 926–929.

55.  Gardner, R.L. and R.G. Edwards, “Control of the sex ratio at full term in the rabbit by 

transferring sexed blastocysts,” Nature, 1968. 218: p. 346–349.

56.  Theodosiou, A.A. and M.H. Johnson, “The politics of human embryo research and 

the motivation to achieve PGD,” Reprod BioMed Online, 2011. 22: p. 457–471.

57.  Edwards, R.G., B.D. Bavister and P.C. Steptoe, “Early stages of fertilization in vitro of 

human oocytes matured in vitro,” Nature, 1969. 221: p. 632–635.

58.  Jones Jr., H.W., “From reproductive immunology to Louise Brown,” Reprod BioMed 

Online, 2002. 4(Suppl.1): p. 6–7.

59.  Edwards, R.G., R.P. Donahue, T.A. Baramki and H.W. Jones Jr., “Preliminary attempts 

to fertlilize human oocytes matured in vitro,” Am J Obstet Gynec, 1966: 96, p. 192–200.

60.  Edwards, R.G., L. Talbert, D. Israelstam, H.N. Nino, and M.H. Johnson, “Diffusion 

chamber for exposing spermatozoa to human uterine secretions,” Am J Obstet Gynec

1968. 102: p. 388–396.

61.  Edwards, R.G., interviewed in: To Mrs. Brown a daughter, 1980. Peter Williams TV: The 

Studio, Boughton, Faversham, UK. 

62.  Anon, “Colin Austin,” Austral Acad Sci Newsletter, 2004. 60: p. 

63.  Chang, M.C., “Fertilizing capacity of spermatozoa deposited into the fallopian tubes,” 

Nature, 1951. 168: p. 697–698. 

64.  Austin, C.R., (1951) “Observations of the penetration of sperm into the mammalian 

egg,” Austral J Sci Res Series B, 1951. 4: p. 581–596.

65. Bavister, 

B.D., “Environmental factors important for in vitro fertilization in the hamster,” 

Reproduction, 1969. 18: p. 544–545.

66.  Edwards, R.G., and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 81–83.

67. “Morris, Prof. Norman Frederick,” In: Who Was Who, A & C Black, 1920–2008: Oxford 

University Press. http://www.ukwhoswho.com/view/article/oupww/whowaswho/


68.  Rock, J. and M. Menkin, “In vitro fertilization and cleavage of human ovarian eggs,” 

Science, 1944. 100: p. 105–107.

69.  Shettles, L.B., “A morula stage of human ovum developed in vitro,” Fert Steril, 1955. 9: 

p. 287–289.

70.  Petrov, G.N., “[Fertilization and first stages of cleavage of human egg in vitro],” 

Arkhiv Anatomii, Gistologii i Embriologii, 1958. 35: p. 88–91.

71.  Yang, W.H., “[The nature of human follicular ova and fertilization in vitro],” J Jap 

Obstet Gynec Soc, 1963. 15: p. 121–130.

72.  Petrucci, D., “Producing transplantable human tissue in the laboratory,” Discovery

1961. 22: p. 278–283.

73.  Hayashi, M., “Fertilization in vitro using human ova,” In: Proceedings of the 7th 

International Planned Parenthood Federation, Singapore, 1963. Excerpta Medica 

International Congress Series No. 72: Amsterdam, Netherlands. pp. 505–510.

74.  “New step towards test-tube babies,” Nature-Times News Service.  The Times, 1969. 

Friday, Feb 14: p. 1.

75.  Steptoe, P.C. and R.G. Edwards, “Laparoscopic recovery of preovulatory human oo-

cytes after priming of ovaries with gonadotrophins,” Lancet, 1970. 295: p. 683–689.

76.  Steptoe P.C., R.G. Edwards and J.M. Purdy, “Human blastocysts grown in culture,” 

Nature, 1971. 229: p. 132–133.

Edwards_lect_.indd   21

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77.  Johnson, M.H., S.B. Franklin, M. Cottingham and N. Hopwood, “Why the Medical 

Research Council refused Robert Edwards and Patrick Steptoe support for research 

on human conception in 1971,” Hum Reprod, 2010. 25: p. 2157–2174.

78.  Edwards, R.G. and P. Steptoe, A Matter of Life: The Story of a Medical Breakthrough. 1980, 

Hutchinson: London, UK. p. 11–15.

79. Pappworth, 


Human Guinea-Pigs, 1967. Routledge and Keegan Paul: London, 


80.  Edwards, R.G. and D.J. Sharpe, “Social values and research in human embryology,” 

Nature, 1971. 231: p. 87–91. 

81.  Edwards, R.G. and P.C. Steptoe, P.C., “Birth after the reimplantation of a human  

embryo,” Lancet, 1978. 312, 366.

82.  Johnson, M.H. and A.A. Theodosiou, “PGD and the making of the genetic embryo 

as a political tool,” In: (Ed. S. McLean) Regulating PGD: A Comparative and Theoretical 

Analysis, 2012. Routledge: London, UK. In the press.

83.  Gardner, R.L. and M.H. Johnson, “Bob Edwards and the first decade of Reproductive 

BioMedicine Online,” In: (Eds. R.L. Gardner, M.H. Johnson) Tenth Anniversary 

Issue 1 Bob Edwards and the first decade of Reproductive BioMedicine Online, 2010. Elsevier: 

Amsterdam, Netherlands. P. 1–24. 

84.  Cohen, J., “From Louise Brown to ESHRE and the journals,” Reprod BioMed Online

2002. 4(Suppl.1): p. 8–10.

85.  Bennett, F., “Birth of a journal: a personal memoire,” Reprod BioMed Online, 2010. 21: 

p. 727–728.

Portrait photo of professor Edwards © Bourn Hall.

Edwards_lect_.indd   22

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