First published in Chemistry in New Zealand in January 2017 and authored by the late Professor Brian Halton.
Much has been written about Rosalind Franklin in books, biographies and web-based items. The Royal Society (London) inaugurated its Franklin award, The Rosalind Franklin University of Medicine and Science in North Chicago, named after her, and college buildings, laboratories and graduate residences are now named in her honour. Despite these, many chemists remain unaware not so much of her crystallographic studies as the environment and attitudes prevalent throughout her short career. It is for this reason that she is the subject of this issue’s Unremembered Chemist. However, to provide balance and perspective, the King’s College DNA studies are briefly summarised.
Rosalind Franklin was born in London on July 25, 1920, the first daughter and second child of Ellis Arthur and Muriel Frances (née Waley) Franklin. Two sons and a second daughter were born after her and David, her elder brother was born in 1919. The family home was in Notting Hill, now an affluent district in North West London but not quite so in the 1920s. Both her parents’ families were involved with and prominent in the London Jewish community. Her great-grandfather Jacob Waley had been a respected barrister and the first Jewish professor in an English University (Political Economics, UC-London) while her uncle Herbert Samuel had been the Home Secretary and became Leader of the Liberal party for four years from 1931. The family was politically involved dominantly in striving for the working classes. Her father was a merchant banker with liberal politics who voluntarily taught at the London Working Men’s (LWM) College (one of the earliest adult education institutions in the UK) throughout his adulthood, on electricity, magnetism and the History of the Great War. He had intended studying physics at Oxford, but was called up to join the army at the outbreak of the First World War instead. His involvement at the LWM had him rise to become Vice-Principal and have the physics laboratory named after him. In the period before WWII Rosalind’s parents helped settle Jewish refugees from Europe and particularly those from the Kindertransport, taking two Jewish children into their own home.
The Franklin family were financially comfortable in their home at 5 Pembridge Place, Notting Hill, reputed to be a lively and happy place with the children an integral part of it. Ellis and Muriel were particularly supportive of their offspring, ensuring that they developed their individuality in comfortable surroundings, treating both sons and daughters with surprising equality given the era in which they lived.1 Rosalind grew up surrounded by brothers, the older David and younger Colin and Roland until the birth of her sister Jenifer nine years later. From them she learned about competition, sports, and other things more typically of interest to boys and is reputed to have often taken to them more than her brothers.2 They were a close family, partaking in lively discussion and vigorous debate at which, according to sister Jenifer, the highly intelligent, logical, determined and articulate Rosalind excelled. Rosalind had such a strong independent streak that she would even argue with her assertive father. She was also unusual in that she ignored dolls, greatly preferring to craft items, draw, photograph, or read, talents she used later in life to sketch, and then build her own molecular models and equipment.2
From early childhood Rosalind Franklin showed exceptional scholarly ability. At the age of six she joined David at the nearby Norland Place (Holland Park) private school where she was reputed3 to ‘do her sums for pleasure’. At age nine, she entered Lindores School for Young Ladies, a boarding school in the seaside town of Bexhill in Sussex; her parents thought it appropriate for her delicate health.3 After two years there and at age eleven she transferred to St. Paul’s Girls’ School in Brook Green, Hammersmith, West London, about 3.5 km from home. It is, and was, a school known for its high standards and success rate, and was one of the few girls’ schools to teach chemistry and physics when Miss Franklin entered. As the Franklin women were expected to focus their education, talents and skills on political, educational, and charitable forms of community service, it was perhaps surprising that Rosalind expressed an early fascination with physics and chemistry classes. At St. Paul’s she excelled in science, Latin and sports, also becoming fluent in French and taking German. She topped her classes, winning annual awards; her sole weakness was in music. This to the extent that her music teacher, the noted Gustav Holst, enquired of her mother if the girl had tonsillitis or suffered from a hearing problem. When she was 15, she decided to become a scientist despite her father being decidedly against ladies using education to gain a career.2,3 His ideal was for Rosalind to gain a diploma and spend her life like her mother. And this was much because career prospects for women in science, and particularly academia, were decidedly poor – no woman held a major post in a British university.1 With six distinctions, a 1938 matriculation, a School Leaving Exhibition scholarship of £30 a year for three years, a scholarship for university, and £5 from her grandfather, Rosalind entered Newnham College at Cambridge a year earlier than many students and against the advice of the Head of the College. Her father asked that she donate the £30 scholarship to a deserving refugee student and told her he wanted to be kept informed of her progress at least weekly.4
On entry to Cambridge University and Newnham College, which with Girton was the only woman’s one, she studied chemistry within the Natural Sciences Tripos graduating in 1941. She was one of the 500 women admitted, a limit set so that the women would not exceed 10% of the total male numbers. These women were not undergraduates but designated students of Girton and Newnham. Oxford University granted women degrees from 1921 but Cambridge steadfastly refused and did not award them a BA Cantab or any other degree deferring, instead, to a ‘degrees titular’. Despite this, the facilities of the university, male supervisors and male research partners were the norm. Marriage was no deterrent to teaching, although such women were not allowed to participate in the affairs of the institution nor allowed to sit with their male colleagues at university ceremonies; instead they were included with the academic wives and without academic regalia. Rosalind’s preliminary (end of first year) examination result in May of 1939 was a first class, a result far above the third class she expected; her self-doubt was strong. The result allowed her to continue to the first part of the Tripos the next year.
Her undergraduate teachers included spectroscopist W.C. Price, with whom she worked while a laboratory demonstrator, and F.S. Dainton, an early pioneer who had recently joined Cambridge to work with Norrish in photochemistry. She attended as many lectures as she could taking in such noted academics as Lawrence Bragg, J.J Thompson and J.B.S. Haldane. Then, in 1940, Adrienne Weill, a French refugee and former student of Marie Curie, arrived at Newnham and became Franklin's mentor and friend. She had a huge influence on Franklin’s life and career. She also helped Rosalind to improve her spoken French. When war broke out in 1939 Rosalind was starting her second year and her father tried, unsuccessfully, to persuade her to abandon her studies for war work ̶ Rosalind took the view that she would be of more use to the war effort if she completed her chemistry degree.2 However, at that time she, and all the other students were expected to continue their study with less supervision because the science staff were occupied elsewhere in the war effort. This suited Rosalind but her independent enthusiasm led her close to the point of exhaustion. Yet she remained sufficiently dutiful to her family that during the university vacations, following the family tradition, she worked with the German-Jewish Refugee Committee in London. With the impact of war very evident by May 1940, Franklin found herself doubting her ability to the extent that she cut labs so as to revise for the first part of the Tripos. Again she expected failure and not the first-class pass she obtained. This gave her a £15 college exhibition scholarship and allowed her to proceed to the final year.
This last year saw her specialising in physical chemistry and attracted to crystallography. She had a supervisor in Newnham College, Delia Simpson, but felt it more appropriate to be under physical chemist and Yorkshire man F.S. (Fred) Dainton. Despite his full schedule, Dainton was sufficiently impressed by Rosalind’s directness, and the assistance she offered other students that he took her on. He is reputed to have told Newnham College that he did not expect Rosalind to gain a first-class degree, not because of inability but because she would spend far too long on one topic rather than moving on to the next and attending to the entire examination paper; apparently she answered two rather than the three questions required on two of her papers.4 She was just as pessimistic in approaching her final exams as those of the first two years. When Franklin graduated from Cambridge in 1941 she (and all women graduands) was not awarded her degree until 1947, when the university changed its regulations. As Dainton had predicted, it was not first class but an upper second, a result that bitterly disappointed her but fitted to his comments, her over-enthusiasm, and her exhaustion. Despite the result, she was offered a scholarship for fourth-year research with Professor Ronald Norrish (the British photochemist and 1967 chemistry Nobel Prize winner). At that time Norrish’s family had moved away from Cambridge to Devon to escape the possibility of bombing and he had succumbed to heavy drinking. This led to a lower than expected security rating and his inability to partake in significant war work. One outcome of this was that he treated his juniors badly and Rosalind Franklin was no exception.4 Her research project was to study further the polymerisation of acetaldehyde and formic acid that had been the subject of a 1935 paper by Norrish and Carruthers.5 Rosalind’s research and study led her to the conclusion that the result anticipated by Norrish was untenable and Dainton agreed with her. This led to a major confrontation between student and professor and eventually a new project. By the summer of 1942 Rosalind Franklin had to decide whether to apply to stay at Cambridge for a PhD under Norrish or leave.4 The decision was in large measure forced upon her by the Ministry of Labour decree that all female research students were to be de-registered and become eligible for military service. Surprisingly perhaps, Norrish encouraged her to apply to retain her reserved status and then painted an unimpressive picture of research in industry. It seems that he had recognised her ability and potential.
When the opportunity of research at the British Coal Utilisation Research Association (BCURA; pronounced B cura) availed itself that 1942 summer Franklin resigned her scholarship, left the Norrish laboratory and began work with the new organisation. The BCURA laboratories were located near Kingston upon Thames and other sites around London, all under the direction of D.H. Bangham.6 He employed young scientists trained in the latest research techniques in his new organisation and, because of the various locations, they had comparative freedom in their work. For the next four years, Rosalind Franklin worked there studying the micro-structures of various coals and carbons. Her research explained why some coals were more permeable to water, gases, and solvents and how heating and carbonisation affected permeability. She found that the pores in coal have fine constrictions at the molecular level and that they increase on heating, depending on the carbon content. They are molecular sieves that successively block the penetration of substances according to molecular size. Franklin was the first to identify and measure the effects of heat on the microstructure of coal which included the changes that occurred during the conversion of some coals to graphite. The work led to a classification of coals and an accurate prediction of their performance, an area in which her expertise clearly was recognised. The results of her study were written-up and submitted for her Cambridge PhD with a thesis, completed back at Newnham College and entitled: The physical chemistry of solid organic colloids with special reference to coal. It was submitted in May 1945 and deposited in the university library the following year.
During her professional role as assistant research officer and graduate student at BCURA, Rosalind shared an uncle’s house on Putney Common with a cousin and her friend, and where her lifestyle became markedly more relaxed. Her work hours were restricted and she took delight in becoming more domesticated, cleaning, cooking and shopping. The four years at BCURA generated five papers, one jointly with Bangham,7 and as sole author of three. Her Cambridge friend and mentor Adrienne Weill had spent the war years working with Lawrence Bragg in Cambridge but returned to Paris as a metallurgist in the French government laboratory for naval research after the 1944 Liberation. Rosalind stayed with BCURA for a year after her PhD and became recognised for her forceful and direct approach to matters in which she knew she was correct, and this irrespective of the place or her position.4 In looking for a post-war position she wrote to Adrienne Weill: If you ever hear of anybody anxious for the services of a physical chemist who knows very little about physical chemistry but a lot about holes in coal, please let me know. As a result, Marcel Mathieu (William Bragg-trained crystallographer working in the Department of Materials in the Ministry of Defence) made contact when he attended the autumn 1946 carbon research conference in London with crystallographer Jaques Méring.1,4 Within weeks Franklin had the offer of a chercheur (postdoctoral) position in Laboratoire Centrale des Services Chemiques de l’Etat (LCSCE) (the ‘labo’) in Paris, downstream from Notre Dame. She accepted.
Rosalind learnt X-ray crystallographic methods and techniques from Méring and immersed herself in her work and French lifestyle. Méring’s expertise was with disordered crystals using low angle monochromatic, highly focussed X-rays, then a technique and speciality of French crystallographers. He was studying graphite. Franklin soon became proficient in X-ray techniques and studied the structure of coal and related carbons. With her skill in preparation she was soon detecting and clarifying the fundamental differences between those carbons that transformed into graphite on heating and those that did not. Her four years at the LCSCE were productive and gave rise to a series of twelve papers from 1950 (Influence in the bonding electrons in the scattering of x-rays by carbon) to 1957 (Changes in the structure of carbon during oxidation) of which only two were co-authored. Soon after her arrival, Vittoria Luzzati (an Italian Jew married to a French medic who had spent the war years in Argentina) joined the ‘labo’ and moved into the room next door; they became strong friends. She worked hard and played hard. Throughout her short life, Rosalind Franklin undertook strenuous hiking, climbing and mountaineering holidays. Initiated when a child by her father in the mountains of Norway, her aptitude for and enjoyment of such vacations was one of the ways she spent time away from the laboratory. Her friendship with Luzzati was cemented by their vigorous debates on science and politics. The photograph below by Luzzati shows her in a cabin in the French Alps.
In France, Rosalind immersed herself in art and culture and made friends with her ‘labo’ colleagues. Despite the short distance from London, travel was not easy (the cross channel ferries had no stabilizers) and her parents expected her to return to London fairly soon. In fact, she applied for a position to work with J.D. Bernal at Birkbeck College (University of London) in the autumn of 1949 and, like Francis Crick (who was leaving the Admiralty in London), she was unsuccessful. She decided to stay in Paris until more of her work was published. Strenuous vacations became the norm with planning as detailed as for a laboratory experiment. Companions, usually female scientists, usually accompanied her.
From early in 1950 Rosalind Franklin began the search for a London position. Her upbringing would not have her take anything in the provinces and so she approached senior academics with whom she had had contact. In particular, she asked Charles Coulson, Head of Theoretical Physics at King's College London and known from her BUCRA time, about ICI research fellowships and the application process. His reply said that were she interested she could apply for one at King’s in the biological application of X-ray techniques. Knowing little biology she responded in the affirmative saying she would learn. Then, in March, she visited Coulson who introduced her to Professor J.T. Randall, the King’s College head of physics and biophysics, the only department in Britain studying biophysics; she applied for a Turner and Newall fellowship in his sphere. Her publications from the Paris studies advanced with the most significant paper appearing in Acta Crystallographica that June.
She holidayed in Italy with Luzzati and his wife, and was then called to London for interview by the Turner and Newall Board early in June. She won a three-year Fellowship to work at King’s under J.T. Randall from the autumn. The departmental secretary advised her that she would be using her X-ray expertise to study proteins in solution and the structural changes involved in them. The salary was set at £800 p.a., almost double that of a Junior Lectureship. As she was reluctant to leave Paris so soon, she applied for deferral until the beginning of 1951 and, with that approved, she went off to Normandy with the Luzzatis.
As time passed, Rosalind became less and less satisfied with her decision to return to London, again doubting herself and feeling that a life in France or Italy would be preferable. Yet change she could not and so she applied her energies to settling the scientific matters concerning her start at King’s College. She suggested improvements for the equipment King’s was to obtain for her and especially in the design of the X-ray camera. All of these went in a late November letter to Randall but his response came as a bombshell. Her entire project was to change. The study of proteins in solution was to become one of certain biological fibres using high and low angle diffraction, an area that now interested him. She was told that the X-ray work would be done by her with PhD student Ray Gosling and Mrs Heller a temporary assistant from Syracuse, New York. The fibres in question were of DNA provided by Professor Signer of Bern to Maurice Wilkins after a Faraday Society lecture in London the previous June and from which Gosling already had good fibre diagrams.1 Whether this change caused Franklin concern or not is not clear but she left the Seine for the Strand at Christmas 1950 more concerned with her relocation home than the work itself.
Following the pre-WWII work of Astbury and Bernal in the UK, the experiments of Avery in New York in the 1940s and the theorising of Erwin Schrödinger in Dublin on life, the scene was set for physicists to advance biology. As Schrödinger put it to explain why genes do not disintegrate as expected from entropy: Life is doing something. Then, in 1949, Chargaff at Columbia College of Physicians and Surgeons found that the number of thymine and cytosine molecules always equalled those of adenine and guanine in DNA, but he could not explain why. And that is where matters stood in January 1951 when Rosalind Franklin entered King’s College London.
Colleagues and friends of Rosalind Franklin considered her to be a brilliant scientist and a kindhearted woman with a personality marred by short temper and stubbornness when she knew she was right. This provided a challenge to those working with her and eventually these included Maurice Wilkins, her colleague at King's College. The contrast of male dominated and bombed London to the intact and liberal Paris were hard for Rosalind to readjust to. King’s was no Cambridge in either its speech or attitudes despite it predating Oxbridge with serious study in laboratory science. Shortly after arrival Rosalind was told that women were not allowed in the King’s senior common room where staff could take lunch, nor was she or any of the MRC women admitted to the upstairs smoking room for coffee afterwards.8 There was, however, a second dining room available to both men and women and some of the male staff preferred to eat there than in the senior common room. This was encouraged by Randall who had many female staff members and liked everyone to get together for morning coffee and afternoon tea and eat lunch as a group there with him. However, the environment was alien to everything Rosalind had become.
On Monday January 8, J.T. Randall held a meeting in his office to introduce Rosalind to her group – Gosling and Heller, and Alec Stokes a physicist and mathematician who was to study theoretical problems. He advised Rosalind that Gosling was now to work with her and not Wilkins.9 Maurice Wilkins, the deputy director who had been working almost exclusively on DNA at King’s for several years and had with Gosling recorded X-ray images, was on holiday. As an aside, Stokes had suggested to Wilkins that what Gosling had shown could result from a helical structure. Now Randall was not simply Wheatstone Professor of Physics at King’s but also head of department and honorary director of the Medical Research Council biophysics unit. He was recognised for appointing highly competent staff and giving prominence to women scientists. He was less successful at settling their responsibilities once he had them. It needs be remembered that by 1951 the Royal Society had elected just seven women Fellows and their number remained below 4% until the end of the 20th century; physics was the most male dominated discipline.
Maurice Wilkins returned from his break some days later unaware that Franklin had been given responsibility for the DNA X-ray work or that Ray Gosling had been transferred to her. She, in turn, did not realise he assumed her to be a member of his team. Nevertheless, the first months proved to be amicable enough with both working separately but cordially on DNA; this to the extent that discussions led to Rosalind being acknowledged in one of his papers,10 then not common from a senior scientist to a female colleague. The pair occasionally lunched together on a Saturday at the Strand Palace Hotel (a good and inexpensive buffet even for academics) and conversation was not difficult as Rosalind had interests in painting, theatre, poetry and existentialism. She was a very different person socially than when at work. Their disagreement stemmed from their conflicting characters ̶ Franklin was forceful especially when she knew her topic and it related to research, whereas Wilkins was shy and retiring and disliked argument. The situation Rosalind found herself in was not improved by her junior status and that, except for Coulson and Randall, her colleagues were unaware of the reputation she had built for herself in the coal area. The situation deteriorated and Gosling found himself very much in the middle and trying to play the peacemaker.9 Eventually, the dispute reached the stage where Professor Randall suggested that Franklin could be better off leaving King’s even though the DNA work was not yet complete. This she did, but not until March 1953 when she joined the Bernal group at Birkbeck on Malet Street.
Rosalind Franklin spent 27 months at King’s College. There Sven Furberg, a Norwegian researcher, had shown from a model he made that the DNA sugars were orthogonal and not parallel to the bases, and stacked like pennies contradictory to Astbury’s prediction. She began her studies using a new fine-focus X-ray tube and micro-camera ordered by Wilkins, but which she refined, adjusted and focused carefully. Working with Gosling, she began to apply her expertise to the structure of DNA. Drawing on her background, she skillfully manipulated the critical hydration of her specimens. Eight months into her new job, in September 1951, she made a pivotal breakthrough by discovering a previously unsuspected second type of DNA that comes from exposure to high levels of moisture. She termed this B DNA. The previously known drier form became A DNA. Furthermore, Rosalind realised that the earlier X-ray studies were less helpful than they might have been because the DNA had contained a mixture of both forms, causing blurring of the photos. She presented this information in a small seminar in King’s in autumn 1951, when Jim Watson was in the audience; he took no notes and was subsequently unable to recall all the information.
On May 2, 1952, Ray Gosling took an X-ray diffraction photo of B DNA that would become both famous and notorious, now termed Photo 51. Rosalind had been able to draw out exceptionally fine DNA fibers and, when appropriately hydrated, were examined using the new equipment that she had designed and ordered. She was then able to quickly establish the crucial differences between the A- and B-forms of DNA. Vittorio Luzzati had suggested that Rosalind incorporate the labour intensive (no computers) Patterson function [Patterson was the British physicist born in Nelson, New Zealand, on July 23, 1902] into her study of DNA while she was on holiday over the 1951 Christmas break. This she did to the X-ray pictures of the A-form and correctly located and measured the positions of the backbone phosphate groups (outside) and the nitrogenous bases (inside) which she hypothesised as being a double stranded helical molecule. She measured the unit cell dimensions, classified the space group as C2, but did not realize that the two sides of the sugar-phosphate backbone ran in opposite directions (anti-parallel). Nonetheless, Photo 51 was clear enough to allow her to determine precisely the 34 Å helical repeat and the 20 Å helical diameter.2 We now know that B-DNA is the usual arrangement within living cells, where the environment is very moist. By January 1953, Franklin had reconciled her conflicting data, concluding that both DNA forms had two helices, and had started to write a series of three draft manuscripts, two of which included a double helical DNA backbone. Her two manuscripts on A-DNA reached Acta Crystallographica on March 6, 1953, one day before Watson and Crick had completed their famous model of B-DNA and before Rosalind knew of their work. On July 8, 1953 she modified one of these, by then in proof items, in light of the King's and Cambridge work. On learning of Watson and Crick’s model, Rosalind rewrote her own draft manuscript on the B molecule as a supportive paper that was published11 in the same April 1953 Nature issue as Watson and Crick12 and that of Wilkins, Stokes and Wilson.13 Franklin had initially found it difficult to interpret her results but had come to the conclusion that DNA had a double helix structure, with component nucleotides or bases on each strand that were complementary, enabling the molecule to replicate. Above all, Franklin noted that an infinite variety of nucleotide sequences would be possible to explain the biological specificity of DNA,2 thereby showing that she had glimpsed the most decisive secret of DNA: the sequence of bases contains the genetic code.
It seems that Rosalind Franklin never knew that her data had been given to Watson and Crick and contributed critically to their proposal. With hindsight, several of Franklin’s new precise findings could have been inferred from formerly published poorer data, but those older data did not sufficiently stimulate Watson and Crick to build a correct model, whereas Franklin’s results did. Furthermore, Franklin distinguished between the A versus B forms of DNA, measured the unit cell dimensions, and identified the space group, which indicated the anti-parallel nature of the backbone. These were new essential pieces of information. What Rosalind Franklin had not known was that on a visit to King’s College on January 30, 1953, Jim Watson was shown (but not given) Photo 51 by Wilkins, and Wilkins a preprint of a Pauling-Corey manuscript14 that included a DNA structure remarkably like their first incorrect model. From the Franklin-Gosling photograph it became obvious to Watson and Crick that that they needed far more than a helix; they needed precise observations from X-ray crystallography and those numbers were unwittingly provided by Franklin herself as they formed part of a brief informal report given to Max Perutz of Cambridge University. In February 1953, Perutz passed the report to Bragg, and he to Watson and Crick.
With those data, Crick had what he needed to perform his calculations. The numbers were decisive in showing the molecule to be in two matching parts running in opposite directions. Franklin’s report was not confidential, and there is no question that the Cambridge duo acquired the data dishonestly. However, they did not tell anyone at King’s what they were doing, and they did not ask Franklin for permission to interpret her data. Whilst undoubtedly cavalier, the behaviour of Perutz, Bragg, Watson and Crick would likely have been just the same had the data come from Maurice Wilkins, and with whom Ray Gosling completed his PhD.
In mid-April 1953, Franklin wrote to Crick from Birkbeck College asking if she could see their model. This she did but still remained sceptical of model building. As an experimental scientist, she was always cautious and wanted significant evidence before publishing anything as proven. This fitted with much of the scientific community hesitating for some years before accepting the double helix proposal.
Rosalind Franklin had decided to leave King’s early in 1952 as the feud with Wilkins had deteriorated to the extent that they were not speaking, and she had taken all the X-ray equipment as was her right according to her appointment letter. She sought a position with Bernal at Birkbeck College, received support and an agreement that she could transfer her Turner and Newell fellowship there. However, Bernal did not have this happen until the following January and, in fact, she did not move until March 1953 so as to complete the DNA work first. Her impending departure signalled to Randall that she might take his school’s work on B-DNA with her and so he got an undertaking from Rosalind that it would remain King’s. This did not mean that she would move to other areas of crystallography rather than stay with biomolecules; she moved her attention from DNA to RNA and the tobacco mosaic virus (TMV). In addition she helped Ray Gosling with his thesis and published the results on the helical nature of A-DNA.15
TMV is a positive-sense single stranded RNA virus that infects a wide range of plants, especially tobacco and other members of the family Solanaceae. It manifests itself as mosaic-like mottling and discoloration on the leaves and was the first virus to be discovered. At Birkbeck with Bernal as Head, Rosalind blossomed, working as a Senior Scientific Officer and team leader with her own research group funded by the Agricultural Research Council. Her work there was as successful as that at King’s and took her through the rest of her short career. She generated a first-rate research team that included subsequent Nobel Laureate Arron Klug, and it provided some sixteen seminal papers on a range of viruses. Klug, a theoretical physicist, chemist and crystallographer, moved to Birkbeck as a Nuffield Fellow in late 1953 on gaining his Cambridge PhD. His office was on the same floor as Rosalind’s and after seeing some of her X-ray images early in 1954 became committed to work with her. The first major work on TMV appeared in 1955 in Nature16 where all the TMV particles were shown to be the same length (3000 Å) in contradiction to the eminent virologist Norman Pirie’s view; he would not accept the result even though subsequently she was proved correct. The disagreement with him meant that from then on she had to grow her own viruses rather than have him provide them. The complete structure of TMV was handled by PhD student Kenneth Holmes and gave a total of eight publications together and with Klug and his student. As a team, the publications on TMV, the cucumber virus and the turnip yellow mosaic virus from 1956 were seminal. By then Rosalind was using the Perutz heavy atom substitution technique of adding electron-dense atoms to the protein without disturbing its structure. Team members continued working on other RNA viruses that affected plants, including the potato, tomato and pea; American postdoctoral Donald Caspar worked on the precise location of RNA molecules in TMV. Complementary papers in the March 10 issue of Nature showed that the RNA in TMV is wound along the inner surface of the hollow virus.17
The ARC research grant was due to expire in 1957 but was extended until March 1958. In applying for the extension, Rosalind sought promotion to Principal Senior Scientific Officer but it was declined. It illustrates the disparity at that time between full-time academic researchers and those who also taught, and between men and women (something the universities began to address in 1955). Rosalind had little choice but to apply for a new grant and got one from the US National Institutes of Health. At £10,000 for three years to work on the polio virus, it was the largest ever received at Birkbeck.4 Then the first major international post-WWII world fair was the held in Brussels in 1958 and Franklin was invited to provide a 1500 mm high model of TMV. She started the build in 1957 using table tennis balls and plastic bicycle handlebar grips and it was displayed in the International Science Pavilion from April 17, one day after she died.
After leaving King’s College and working on TMV, Rosalind became friendly with both Watson and Crick, though more so with Crick and his wife. She spent time at their home4 and toured Spain with them in the spring of 1956. Francis Crick noticed changes in Rosalind’s health but with her high reserve he did not pursue the topic with her. A little later that year, while on one of a few work-related trips to the US, she began to suspect a health problem and when back in London she sought medical advice. The outcome was an operation on September 4 that revealed two tumours in her abdomen. Following the surgery and other periods of hospitalisation, Rosalind spent time convalescing with various friends and family members that included the Cricks. She continued working throughout the following two years, despite having three operations and experimental chemotherapy. She experienced a 10-month remission and worked up until several weeks before her death. Even while undergoing cancer treatment, Rosalind continued to work, and her group produced seven more papers in 1956 and six in 1957. At the end of 1957, she again fell ill and was admitted to the Royal Marsden Hospital. She made her will at the beginning of December naming Aron Klug principal beneficiary, then returned to the lab in January 1958. She was promoted to Research Associate in Biophysics on February 25 but fell ill again on March 30, and died on April 16, 1958, in Chelsea, London, of bronchopneumonia, secondary carcinomatosis (widespread dissemination of carcinoma), and ovarian cancer.
Exposure to X-ray radiation could be a factor in her illness as safety precautions were not as stringent in her time as they are now, and it was known that she had exceeded the limit of her dosimeter in France. However, her own DNA may have predisposed her to ovarian cancer as it is known that Ashkenazi Jews, who settled and established communities throughout Central and Eastern Europe, have a predisposition to it. Moreover, other members of her family had died of gynaecological cancer. Undoubtedly, Rosalind Franklin was brave through the final stages of her cancer. Unable to walk, she crawled up stairways between laboratories at Birkbeck insisting that she continue to work. Her Birkbeck research team comprising Klug, Finch and Holmes moved to the Laboratory of Molecular Biology in Cambridge in 1962.
Her death of ovarian cancer on April 16, 1958 was four years before the Nobel Prize was awarded to Watson, Crick and Wilkins for their DNA work. She never learned the full extent to which Watson and Crick had relied on her data to make their model; if she suspected, she did not express any bitterness or frustration. She could not have received a share in the award unless she had been nominated before or in the year of her death. She is buried in the Franklin family plot at the United Jewish Cemetery in Willesden, London.18 The British Heritage foundation attached a plaque to the outside of her apartment at Donovan Court, 107 Drayton Gardens, Chelsea, in 1992.
References and Notes
1. Miksic, M.C., Rosalind Elsie Franklin. In Women in Chemistry and Physics (Grinstein, L.S.; Rose, R.K.; Rafailovich, M.H. eds.), Greenwood Press, Westport, CN, 1993, 191-200.
2. Elkin, L. Rosalind Elsie Franklin. In Jewish Women’s Archive: A Comprehensive Historical Encyclopedia, 2009; see Jewish Women's Archive at: http://jwa.org/encyclopedia/article/franklin-rosalind (accessed 01/09/2016).
3 The biotechnologists. In What is Biotechnology, a collection of information and resources about some of the people whose work has helped build biotechnology into one of the most important tools in our lives today; see: http://www.whatisbiotechnology.org/people/Franklin (accessed 01/09/2016).
4. Maddox, B. Rosalind Franklin the dark lady of DNA, Harper-Collins, New York 2002, pp. xix, 380.
5. Carruthers, J.E.; Norrish, R.G.W. The polymerisation of gaseous formaldehyde and acetaldehyde, Trans. Faraday Soc., 1936, 32, 195-208.
6. Rosalind Franklin: A Crucial Contribution, Scitable - Nature Education; see: http://www.nature.com/scitable/topicpage/rosalind-franklin-a-crucial-contribution-6538012 (accessed 06/09/2016).
7 Bangham, D.H.; Franklin, R.E. Thermal expansion of coals and carbonised coals, Trans. Faraday Soc. 1946, 42B, 289-294.
8. Elkin, L.O. Rosalind Franklin and the double helix, Physics Today 2003, 56(3), 42-48.
9. Attar, N. Raymond Gosling: the man who crystallized genes, Genome Biol. 2013, 14, 402-413.
10. Wilkins, M.H.F.; Gosling, R.G.; Seeds, W.E. Physical studies of nucleic acid, Nature 1951, 167, 759-760.
11. Franklin, R.E.; Gosling, R.G. Molecular configuration in sodium thymonucleate, Nature 1953, 171, 740-741.
12. Watson, J. D.; Crick, F. H. C. Molecular structure of nucleic acids. A structure for deoxyribose nucleic acid, Nature, 1953, 171, 737-8.
13. Wilkins, M.H.F.; Stokes, A.R.; Wilson, H.R. Molecular structure of deoxypentose nucleic acids, Nature, 1953, 171,738-740.
14. Pauling, L.; Corey, R. A proposed structure for the nucleic acids, Proc. Natl. Acad. Sci. 1953, 39, 84-97.
15. Franklin, RE; Gosling, RG. Evidence for 2-chain helix in crystalline structure of sodium deoxyribonucleate, Nature. 1953, 172, 156–157.
16. Franklin, RE. Structure of Tobacco Mosaic Virus, Nature, 1955, 175, 379–381.
17 Caspar, D.L.D. Radial density distribution in the tobacco mosaic virus particle, Nature 1956, 177, 928; Franklin, R.E. Location of the ribonucleic acid in the tobacco mosaic virus particle, Nature 1956, 177, 928-930.
18. Rosalind Franklin at Find a Grave (findagrave.com). The photograph was placed there by Julia Keld (46812479); see: http://www.findagrave.com/cgi-bin/fg.cgi?page=gr&GRid=5858699 (accessed 16/09/2016).
Other books on Rosalind Franklin:
Glynn J. My sister Rosalind Franklin, Oxford University Press 2012, pp. 172.
Sayre, A. Rosalind Franklin and DNA, Norton, New York, 1975, pp. 221; paperback, Norton, 2000, pp. 240.
