Rosalind Franklin

Franklin went to Newnham College, Cambridge in 1938 and studied chemistry within the Natural Sciences Tripos. There she met the spectroscopist Bill Price, who worked with her as a laboratory demonstrator and who later became one of her senior colleagues at King's College London.[32] In 1941, she was awarded second-class honours from her final exams. The distinction was accepted as a bachelor's degree in qualifications for employment. Cambridge began awarding titular B.A. and M.A. degrees to women from 1947, and the previous women graduates retroactively received these.[33] In her last year at Cambridge, she met a French refugee Adrienne Weill, a former student of Marie Curie, who had a huge influence on her life and career and helped her to improve her spoken French.[34][35]

Franklin was awarded a research fellowship at Newnham College, with which she joined the physical chemistry laboratory of the University of Cambridge to work under Ronald George Wreyford Norrish, who later won the Nobel Prize in Chemistry. In her one year of work there, she did not have much success.[36] As described by his biographer, Norrish was "obstinate and almost perverse in argument, overbearing and sensitive to criticism".[37] He could not decide for her what to work upon, and at that time was succumbing to heavy drinking. Franklin wrote that he made her despise him completely.[38] Resigning from Norrish's Lab, she fulfilled the requirements of the National Service Acts by working as an assistant research officer at the British Coal Utilisation Research Association (BCURA) in 1942.[9] The BCURA was located on the Coombe Springs Estate near Kingston upon Thames near the southwestern boundary of London. Norrish acted as advisor to the military at BCURA. John G. Bennett was the director. Marcello Pirani and Victor Goldschmidt, both refugees from the Nazis, were consultants and lectured at BCURA while Franklin worked there.[2] During her BCURA research, she stayed at Adrienne Weill's boarding house in Cambridge until her cousin Irene Franklin asked to join her in a vacated house of her uncle in Putney. With Irene, she volunteered as an Air Raid Warden and regularly made patrols to see the welfare of people during air raids.[39]

She studied the porosity of coal using helium to determine its density.[40] Through this, she discovered the relationship between the fine constrictions in the pores of coals and the permeability of the porous space. By concluding that substances were expelled in order of molecular size as temperature increased, she helped classify coals and accurately predict their performance for fuel purposes and for production of wartime devices such as gas masks.[41] This work was the basis of her Ph.D. thesis The physical chemistry of solid organic colloids with special reference to coal for which the University of Cambridge awarded her a Ph.D. in 1945.[4] It was also the basis of several papers.[2]

With World War II ending in 1945, Franklin asked Adrienne Weill for help and to let her know of job openings for "a physical chemist who knows very little physical chemistry, but quite a lot about the holes in coal". At a conference in the autumn of 1946, Weill introduced her to Marcel Mathieu, a director of the Centre national de la recherche scientifique (CNRS), the network of institutes that comprise the major part of the scientific research laboratories supported by the French government. This led to her appointment with Jacques Mering at the Laboratoire Central des Services Chimiques de l'État in Paris. She joined the labo (as referred to by the staff) of Mering on 14 February 1947 as one of the fifteen chercheurs (researchers).[42][43]

Mering was an X-ray crystallographer who applied X-ray diffraction to the study of rayon and other amorphous substances, in contrast to the thousands of regular crystals that had been studied by this method for many years.[2] He taught her the practical aspects of applying X-ray crystallography to amorphous substances. This presented new challenges in the conduct of experiments and the interpretation of results. Franklin applied them to further problems related to coal and to other carbonaceous materials, in particular the changes to the arrangement of atoms when these are converted to graphite.[2] She published several further papers on this work which has become part of the mainstream of the physics and chemistry of coal and carbon. She coined the terms graphitising and non-graphitising carbon. The coal work was covered in a 1993 monograph,[44] and in the regularly-published textbook Chemistry and Physics of Carbon.[45] Mering continued the study of carbon in various forms, using X-ray diffraction and other methods.[46]

Double helix William Astbury Oswald Avery Florence Bell Lawrence Bragg Erwin Chargaff Francis Crick Michael Creeth Jerry Donohue Rosalind Franklin Raymond Gosling Frederick Griffith John Masson Gulland Denis Jordan Phoebus Levene Friedrich Miescher Fred Neufeld Sir John Randall Alex Stokes James Watson Maurice Wilkins Herbert Wilson Photo 51

In 1950, Franklin was granted a three-year Turner & Newall Fellowship to work at King's College London. In January 1951, she started working as a research associate in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall.[47] She was originally appointed to work on X-ray diffraction of proteins and lipids in solution, but Randall redirected her work to DNA fibres[48] because of new developments in the field, and she was to be the only experienced experimental diffraction researcher at King's at the time.[49][50] Randall made this reassignment, even before Franklin started working at King's, because of the pioneering work by DNA researcher Maurice Wilkins, and he reassigned Raymond Gosling, the graduate student who had been working with Wilkins, to be her assistant.[51]

Even using crude equipment, Wilkins and Gosling had obtained an outstanding diffraction picture of DNA which sparked further interest in this molecule.[52] They had been carrying out X-ray diffraction analysis of DNA in the unit since May 1950, but Randall had not informed them that he had asked Franklin to take over both the DNA diffraction work and guidance of Gosling's thesis.[53] Randall's lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.[54]

Franklin, now working with Gosling,[55] started to apply her expertise in X-ray diffraction techniques to the structure of DNA. She used a new fine-focus X-ray tube and microcamera ordered by Wilkins, but which she refined, adjusted and focused carefully. Drawing upon her physical chemistry background, she also skillfully manipulated the critical hydration of her specimens.[56] When Wilkins inquired about this improved technique, she replied in terms which offended Wilkins as Franklin had "an air of cool superiority".[57]

Franklin presented their data at a lecture in November 1951, in King's College London. In her lecture notes, Franklin wrote the following:[58]

"The results suggest a helical structure (which must be very closely packed) containing 2, 3 or 4 co‐axial nucleic acid chains per helical unit, and having the phosphate groups near the outside."

Franklin's habit of intensely looking people in the eye while being concise, impatient and direct unnerved many of her colleagues. In stark contrast, Wilkins was very shy, and slowly calculating in speech while he avoided looking anyone directly in the eye.[59] In spite of the intense atmosphere, Franklin and Gosling discovered that there were two forms of DNA: at high humidity (when wet) the DNA fibre became long and thin; when it was dried it became short and fat.[60][61]

Franklin named these two forms "B" and "A" respectively. (The biological functions of A-DNA were only discovered 60 years later.[62]) Because of the intense personality conflict developing between Franklin and Wilkins, Randall[63] divided the work on DNA. Franklin chose the data rich "A" form while Wilkins selected the "B" form[63][64] because, according to his self-written biography, Wilkins' preliminary pictures had hinted it might be helical. The X-ray diffraction pictures, including the landmark Photo 51 taken by Franklin's student Gosling at this time,[52] have been called by John Desmond Bernal as "amongst the most beautiful X-ray photographs of any substance ever taken".[60]

By the end of 1951 it was generally accepted at King's that the B form of DNA was a helix, but after she had recorded an asymmetrical image in May 1952, Franklin became unconvinced that the A form of DNA was a helix.[65] In July 1952, as a practical joke on Wilkins (who frequently expressed his view that both forms of DNA were helical), Franklin and Gosling produced a funeral notice regretting the 'death' of helical crystalline DNA (A-DNA).[66] During 1952, they worked at applying the Patterson function to the X-ray pictures of DNA they had produced.[67] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.[68][69]

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 (see below). Her two A-DNA manuscripts reached Acta Crystallographica in Copenhagen on 6 March 1953, one day before Crick and Watson had completed their model on B-DNA. She must have mailed them while the Cambridge team was building their model, and certainly had written them before she knew of their work.[70] On 8 July 1953 she modified one of these "in proof" Acta articles, "in light of recent work" by the King's and Cambridge research teams.[71]

The third draft paper was on the B form of DNA, dated 17 March 1953, which was discovered years later amongst her papers,[72] by Franklin's Birkbeck colleague, Aaron Klug. He then published an evaluation of the draft's close correlation with the third of the original trio of 25 April 1953 Nature DNA articles.[73] Klug designed this paper to complement the first article he had written defending Franklin's significant contribution to DNA structure.[74] He had written this first article in response to the incomplete picture of Franklin's work depicted in Watson's 1968 memoir, The Double Helix.

As vividly described in The Double Helix, on 30 January 1953, Watson travelled to King's carrying a preprint of Linus Pauling's incorrect proposal for DNA structure. Since Wilkins was not in his office, Watson went to Franklin's lab with his urgent message that they should all collaborate before Pauling discovered his error. The unimpressed Franklin became angry when Watson suggested she did not know how to interpret her own data. Watson hastily retreated, backing into Wilkins who had been attracted by the commotion. Wilkins commiserated with his harried friend and then showed Watson Franklin's DNA X-ray image.[75] Watson, in turn, showed Wilkins a prepublication manuscript by Pauling and Corey, which contained a DNA structure remarkably like their first incorrect model.[76]

In February 1953, James Watson and Francis Crick of the Cavendish Laboratory in Cambridge University had started to build a molecular model of the B form of DNA using data similar to that available to both teams at King's. Much of their data was derived directly from research done at King's by Wilkins and Franklin. Franklin's research was completed by February 1953, ahead of her move to Birkbeck, and her data was critical.[77] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,[64][78] but Franklin was opposed to prematurely building theoretical models, until sufficient data were obtained to properly guide the model building. She took the view that building a model was to be undertaken only after enough of the structure was known.[65][79]

Ever cautious, she wanted to eliminate misleading possibilities. Photographs of her Birkbeck work table show that she routinely used small molecular models, although certainly not ones on the grand scale successfully used at Cambridge for DNA. In the middle of February 1953, Crick's thesis advisor, Max Perutz, gave Crick a copy of a report written for a Medical Research Council biophysics committee visit to King's in December 1952, containing many of Franklin's crystallographic calculations.[80]

Since Franklin had decided to transfer to Birkbeck College and Randall had insisted that all DNA work must stay at King's, Wilkins was given copies of Franklin's diffraction photographs by Gosling. By 28 February 1953, Watson and Crick felt they had solved the problem enough for Crick to proclaim (in the local pub) that they had "found the secret of life".[81] However, they knew they must complete their model before they could be certain.[82]

Watson and Crick finished building their model on 7 March 1953, one day before they received a letter from Wilkins stating that Franklin was finally leaving and they could put "all hands to the pump".[83] This was also one day after Franklin's two A-DNA papers had reached Acta Crystallographica. Wilkins came to see the model the following week, according to Franklin's biographer Brenda Maddox on 12 March, and allegedly informed Gosling on his return to King's.[84]

It is uncertain how long it took for Gosling to inform Franklin at Birkbeck, but her original 17 March B-DNA manuscript does not reflect any knowledge of the Cambridge model. Franklin did modify this draft later before publishing it as the third in the trio of 25 April 1953 Nature articles. On 18 March,[85] in response to receiving a copy of their preliminary manuscript, Wilkins penned the following: "I think you're a couple of old rogues, but you may well have something".[86]

Weeks later, on 10 April, Franklin wrote to Crick for permission to see their model.[87] Franklin retained her scepticism for premature model building even after seeing the Watson–Crick model, and remained unimpressed. She is reported to have commented, "It's very pretty, but how are they going to prove it?" As an experimental scientist, Franklin seems to have been interested in producing far greater evidence before publishing-as-proven a proposed model. As such, her response to the Watson–Crick model was in keeping with her cautious approach to science.[88]

Crick and Watson then published their model in Nature on 25 April 1953, in an article describing the double-helical structure of DNA with only a footnote acknowledging "having been stimulated by a general knowledge of" Franklin and Wilkins' "unpublished" contribution.[89] Actually, although it was the bare minimum, they had just enough specific knowledge of Franklin and Gosling's data upon which to base their model. As a result of a deal struck by the two laboratory directors, articles by Wilkins and Franklin, which included their X-ray diffraction data, were modified and then published second and third in the same issue of Nature, seemingly only in support of the Crick and Watson theoretical paper which proposed a model for the B form of DNA.[90][91] Most of the scientific community hesitated several years before accepting the double helix proposal. At first mainly geneticists embraced the model because of its obvious genetic implications.[92][93][94]

An electron micrograph of tobacco mosaic virus

Franklin left King's College London in mid-March 1953 for Birkbeck College, in a move that had been planned for some time and that she described (in a letter to Adrienne Weill in Paris) as "moving from a palace to the slums ... but pleasanter all the same."[95] She was recruited by physics department chair John Desmond Bernal,[96] a crystallographer who was a communist, known for promoting women crystallographers. Her new laboratories were housed in 21 Torrington Square, one of a pair of dilapidated and cramped Georgian houses containing several different departments; Franklin frequently took Bernal to task over the careless attitudes of some of the other laboratory staff, notably after workers in the pharmacy department flooded her first-floor laboratory with water on one occasion.[97]

Despite the parting words of Bernal to stop her interest in nucleic acids, she helped Gosling to finish his thesis, although she was no longer his official supervisor. Together they published the first evidence of double helix in the A form of DNA in the 25 July issue of Nature.[98] At the end of 1954, Bernal secured funding for Franklin from the Agricultural Research Council (ARC), which enabled her to work as a senior scientist supervising her own research group.[99][100] John Finch, a physics student from King's College London, subsequently joined Franklin's group, followed by Kenneth Holmes, a Cambridge graduate, in July 1955. Despite the ARC funding, Franklin wrote to Bernal that the existing facilities remained highly unsuited for conducting research "...my desk and lab are on the fourth floor, my X-ray tube in the basement, and I am responsible for the work of four people distributed over the basement, first and second floors on two different staircases."[100]

Franklin continued to explore another major nucleic acid, RNA, a molecule equally central to life as DNA. She again used X-ray crystallography to study the structure of the tobacco mosaic virus (TMV), an RNA virus. Her meeting with Aaron Klug in early 1954 led to a longstanding and successful collaboration. Klug had just then earned his PhD from Trinity College, Cambridge, and joined Birkbeck in late 1953. In 1955 Franklin published her first major works on TMV in Nature, in which she described that all TMV virus particles were of the same length.[101] This was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct.[102]

Franklin assigned the study of the complete structure of TMV to her PhD student Holmes. They soon discovered (published in 1956) that the covering of TMV was protein molecules arranged in helices.[103] Her colleague Klug worked on spherical viruses with his student Finch, with Franklin coordinating and overseeing the work.[104] As a team, from 1956 they started publishing seminal works on TMV,[105] cucumber virus 4 and turnip yellow mosaic virus.[106]

Franklin also had a research assistant, James Watt, subsidised by the National Coal Board and was now the leader of the ARC group at Birkbeck.[107] The Birkbeck team members continued working on RNA viruses affecting several plants, including potato, turnip, tomato and pea.[108] In 1955 the team was joined by an American post-doctoral student Donald Caspar. He worked on the precise location of RNA molecules in TMV. In 1956 he and Franklin published individual but complementary papers in the 10 March issue of Nature, in which they showed that the RNA in TMV is wound along the inner surface of the hollow virus.[109][110] Caspar was not an enthusiastic writer, and Franklin had to write the entire manuscript for him.[111]

In 1957 her research grant from ARC expired, and was given a one-year extension ending in March 1958.

Expo 58, the first major international fair after World War II, was to be held in Brussels in 1958.[112][113] Franklin was invited to make a five-foot high model of TMV, which she started in 1957. Her materials included table tennis balls and plastic bicycle handlebar grips.[114] The Brussels world's fair, with an exhibit of her virus model at the International Science Pavilion, opened on 17 April, one day after she died.[115]

In 1956, Franklin visited the University of California, Berkeley, where colleagues had suggested her group research the polio virus.[116] In 1957 she applied for a grant from the United States Public Health Service of the National Institutes of Health, which approved £10,000 for three years, the largest fund ever received at Birkbeck.[117][118] In her grant application, Franklin mentioned her new interest in animal virus research. She obtained Bernal's consent in July 1957, though serious concerns were raised after she disclosed her intentions to research live, instead of killed, polio virus at Birkbeck. Eventually, Bernal arranged for the virus to be safely stored at the London School of Hygiene and Tropical Medicine during the group's research. With her group, Franklin then commenced deciphering the structure of the polio virus while it was in a crystalline state. She attempted to mount the virus crystals in capillary tubes for X-ray studies, but was forced to end her work due to her rapidly failing health.[119]

After Franklin's death, Klug succeeded her as group leader, and he, Finch and Holmes continued researching the structure of the polio virus. They eventually succeeded in obtaining extremely detailed X-ray images of the virus. In June 1959, Klug and Finch published the group's findings, revealing the polio virus to have icosahedral symmetry, and in the same paper suggested the possibility for all spherical viruses to possess the same symmetry, as it permitted the greatest possible number (60) of identical structural units.[120] The team moved to the Laboratory of Molecular Biology, Cambridge in 1962,[121] and the old Torrington Square laboratories were demolished four years later, in May 1966.[122]

In mid-1956, while on a work-related trip to the United States, Franklin first began to suspect a health problem. While in New York she found difficulty in zipping her skirt; her stomach had bulged. Back in London she consulted Mair Livingstone, who asked her, "You're not pregnant?" to which she retorted, "I wish I were." Her case was marked "URGENT".[143] An operation on 4 September of the same year revealed two tumours in her abdomen.[144] After this period and other periods of hospitalization, Franklin spent time convalescing with various friends and family members. These included Anne Sayre, Francis Crick, his wife Odile, with whom Franklin had formed a strong friendship,[137] and finally with the Roland and Nina Franklin family where Rosalind's nieces and nephews bolstered her spirits.

Franklin chose not to stay with her parents because her mother's uncontrollable grief and crying upset her too much. Even while undergoing cancer treatment, Franklin continued to work, and her group continued to produce results – seven papers in 1956 and six more in 1957.[145] At the end of 1957, Franklin again fell ill and she was admitted to the Royal Marsden Hospital. On 2 December, she made her will. She named her three brothers as executors and made her colleague Aaron Klug the principal beneficiary, who would receive £3,000 and her Austin car. Her other friends Mair Livingstone would get £2,000, Anne Piper £1,000, and her nurse Miss Griffith £250. The remainder of the estate was to be used for charities.[146]

She returned to work in January 1958, and she was given a promotion to Research Associate in Biophysics on 25 February.[147] She fell ill again on 30 March, and she died on 16 April 1958, in Chelsea, London,[148][149] of bronchopneumonia, secondary carcinomatosis, and ovarian cancer. Exposure to X-ray radiation is sometimes considered to be a possible factor in her illness.[150]

Other members of her family have died of cancer, and the incidence of gynaecological cancer is known to be disproportionately high among Ashkenazi Jews.[151] Her death certificate states: A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.[152] She was interred on 17 April 1958 in the family plot at Willesden United Synagogue Cemetery at Beaconsfield Road in London Borough of Brent. The inscription on her tombstone reads:[153][154]

IN MEMORY OF
ROSALIND ELSIE FRANKLIN
מ' רחל בת ר' יהודה [Rochel/Rachel daughter of Yehuda, her Hebrew name]
DEARLY LOVED ELDER DAUGHTER OF
ELLIS AND MURIEL FRANKLIN
25TH JULY 1920 – 16TH APRIL 1958
SCIENTIST
HER RESEARCH AND DISCOVERIES ON
VIRUSES REMAIN OF LASTING BENEFIT
TO MANKIND
ת נ צ ב ה [Hebrew initials for "her soul shall be bound in the bundle of life"]

Anne Sayre, Franklin's friend and one of her biographers, stated in her book Rosalind Franklin and DNA, "In 1951 ... King's College London as an institution, was not distinguished for the welcome that it offered to women ... Rosalind ... was unused to purdah [a religious and social institution of female seclusion] ... there was one other woman scientist on the laboratory staff".[155] Andrzej Stasiak states "Sayre's book became widely cited in feminist circles for exposing rampant sexism in science."[156] Farooq Hussain states "there were seven women in the biophysics department ... Jean Hanson became an FRS, Dame Honor B. Fell, Director of Strangeways Laboratory, supervised the biologists".[157] Maddox states, "Randall ... did have many women on his staff ... they found him ... sympathetic and helpful."[158]

Sayre states "that while the male staff at King's lunched in a large, comfortable, rather clubby dining room" the female staff of all ranks "lunched in the student's hall or away from the premises".[159][160] Elkin states that most of the MRC group typically ate lunch together (including Franklin) in the mixed dining room discussed below.[59] And Maddox states, of Randall, "He liked to see his flock, men and women, come together for morning coffee, and at lunch in the joint dining room, where he ate with them nearly every day."[158] Francis Crick also commented that "her colleagues treated men and women scientists alike."[161]

Sayre also discusses at length Franklin's struggle in pursuing science, particularly her father's concern about women in academic professions.[162] This account had been taken to accuse Ellis Franklin of sexism against his daughter. A good deal of information explicitly claims that he strongly opposed her entering Newnham College.[163][164][165][166] Franklin's Public Broadcasting Service (PBS) biography goes further by stating that he refused to pay her fees, and that an aunt stepped in for her.[167] Her sister Jenifer Glynn claimed that these stories are myths, and that her parents fully supported Franklin's entire career.[168]

Sexism is said to pervade the memoir of one peer, James Watson, in his book The Double Helix published 10 years after Franklin's death and after Watson had returned from Cambridge to Harvard.[169] His Cambridge colleague, Peter Pauling, wrote in a letter, "Morris [sic] Wilkins is supposed to be doing this work; Miss Franklin is evidently a fool."[170] Crick acknowledges later, "I'm afraid we always used to adopt – let's say, a patronizing attitude towards her."[171]

Glynn accuses Sayre of making her sister a feminist heroine,[172] and Watson's The Double Helix as the root of what she calls "Rosalind Industry". She conjectures that these alleged sexism stories would "have embarrassed her [Rosalind Franklin] almost as much as Watson's account would have upset her",[168] and declared that "she was never a feminist."[173] Klug and Crick also concurred that she was definitely not a feminist.[174]

Franklin's letter to her parents in January 1939 is often taken as reflecting her own prejudiced attitude and that she was "not immune to the sexism rampant in these circles." In it she remarked one lecturer as "very good, though female."[175] But as Maddox explains, it was more of circumstantial comment rather than a gender bias. It was more of an admiration because at the time woman teachers of science were a rarity. She in fact laughed at men who were embarrassed by the appointment of the first female professor Dorothy Garrod.[176]

Rosalind Franklin's first important contributions to the model popularised by Crick and Watson was her lecture at the seminar in November 1951, where she presented to those present, among them Watson, the two forms of the molecule, type A and type B, her position being that the phosphate units are located in the external part of the molecule. She also specified the amount of water to be found in the molecule in accordance with other parts of it, data that have considerable importance in terms of the stability of the molecule. Franklin was the first to discover and formulate these facts, which in fact constituted the basis for all later attempts to build a model of the molecule. However, Watson, at the time ignorant of the chemistry, failed to comprehend the crucial information, and this led to construction of a wrong model.[177]

The other contribution included an X-ray photograph of B-DNA (called Photo 51)[178] taken by Franklin's student Gosling that was briefly shown to Watson by Wilkins in January 1953,[179][180] and a report written for an MRC biophysics committee visit to King's in December 1952 which was shown by Perutz at the Cavendish Laboratory to both Crick and Watson. This MRC report contained data from the King's group, including some of Franklin's and Gosling's work, and was given to Crick – who was working on his thesis on haemoglobin structure – by his thesis supervisor Perutz, a member of the visiting committee.[181][182]

Sayre's biography of Franklin contains a story[183] alleging that the photograph 51 in question was shown to Watson by Wilkins without Franklin's permission,[156][184][185] and that this constituted a case of bad science ethics.[186] Others dispute this story, asserting that Wilkins had been given photograph 51 by Franklin's Ph.D. student Gosling because she was leaving King's to work at Birkbeck, and there was allegedly nothing untoward in this transfer of data to Wilkins[179][187] because Director Randall had insisted that all DNA work belonged exclusively to King's and had instructed Franklin in a letter to even stop working on it and submit her data.[188] Also, it was implied by Horace Freeland Judson, that Maurice Wilkins had taken the photograph out of Franklin's drawer, but this is also said to be incorrect.[189]

Likewise, Perutz saw "no harm" in showing an MRC report containing the conclusions of Franklin and Gosling's X-ray data analysis to Crick, since it had not been marked as confidential, although "The report was not expected to reach outside eyes".[190] Indeed, after the publication of Watson's The Double Helix exposed Perutz's act, he received so many letters questioning his judgment that he felt the need to both answer them all[191] and to post a general statement in Science excusing himself on the basis of being "inexperienced and casual in administrative matters".[192]

Perutz also claimed that the MRC information was already made available to the Cambridge team when Watson had attended Franklin's seminar in November 1951. A preliminary version of much of the important material contained in the 1952 December MRC report had been presented by Franklin in a talk she had given in November 1951, which Watson had attended but not understood.[70][193]

The Perutz letter was as said one of three letters, published with letters by Wilkins and Watson, which discussed their various contributions. Watson clarified the importance of the data obtained from the MRC report as he had not recorded these data while attending Franklin's lecture in 1951. The upshot of all this was that when Crick and Watson started to build their model in February 1953 they were working with critical parameters that had been determined by Franklin in 1951, and which she and Gosling had significantly refined in 1952, as well as with published data and other very similar data to those available at King's. It was generally believed that Franklin was never aware that her work had been used during construction of the model,[194] but Gosling asserted in his 2013 interview that, "Yes. Oh, she did know about that."[195]

Upon the completion of their model, Crick and Watson had invited Wilkins to be a co-author of their paper describing the structure.[196] Wilkins turned down this offer, as he had taken no part in building the model.[197] He later expressed regret that greater discussion of co-authorship had not taken place as this might have helped to clarify the contribution the work at King's had made to the discovery.[198] There is no doubt that Franklin's experimental data were used by Crick and Watson to build their model of DNA in 1953. Some, including Maddox, have explained this citation omission by suggesting that it may be a question of circumstance, because it would have been very difficult to cite the unpublished work from the MRC report they had seen.[84]

Indeed, a clear timely acknowledgment would have been awkward, given the unorthodox manner in which data were transferred from King's to Cambridge. However, methods were available. Watson and Crick could have cited the MRC report as a personal communication or else cited the Acta articles in press, or most easily, the third Nature paper that they knew was in press. One of the most important accomplishments of Maddox's widely acclaimed biography is that Maddox made a well-received case for inadequate acknowledgement. "Such acknowledgement as they gave her was very muted and always coupled with the name of Wilkins".[199]

Fifteen years after the fact, the first clear recitation of Franklin's contribution appeared as it permeated Watson's account, The Double Helix, although it was buried under descriptions of Watson's (often quite negative) regard towards Franklin during the period of their work on DNA. This attitude is epitomized in the confrontation between Watson and Franklin over a preprint of Pauling's mistaken DNA manuscript.[200] Watson's words impelled Sayre to write her rebuttal, in which the entire chapter nine, "Winner Take All" has the structure of a legal brief dissecting and analyzing the topic of acknowledgement.[201]

Sayre's early analysis was often ignored because of perceived feminist overtones in her book. Watson and Crick did not cite the X-ray diffraction work of Wilkins and Franklin in their original paper, though they admit having "been stimulated by a knowledge of the general nature of the unpublished experimental results and ideas of Dr. M. H. F. Wilkins, Dr. R. E. Franklin and their co-workers at King's College London".[89] In fact, Watson and Crick cited no experimental data at all in support of their model. Franklin and Gosling's publication of the DNA X-ray image, in the same issue of Nature, served as the principal evidence:

"Thus our general ideas are not inconsistent with the model proposed by Watson and Crick in the preceding communication".[202]

Franklin was never nominated for a Nobel Prize.[203][204] Her work was a crucial part in the discovery of DNA's structure, which along with subsequent related work led to Francis Crick, James Watson, and Maurice Wilkins being awarded a Nobel Prize in 1962.[205] She had died in 1958, and during her lifetime the DNA structure was not considered to be fully proven. It took Wilkins and his colleagues about seven years to collect enough data to prove and refine the proposed DNA structure. Moreover, its biological significance, as proposed by Watson and Crick, was not established. General acceptance for the DNA double helix and its function did not start until late in the 1950s, leading to Nobel nominations in 1960, 1961, and 1962 for Nobel Prize in Physiology or Medicine, and in 1962 for Nobel Prize in Chemistry.[206] The first breakthrough was from Matthew Meselson and Franklin Stahl in 1958, who experimentally showed the DNA replication of a bacterium Escherichia coli.[207] Now known as Meselson–Stahl experiment, DNA was found to replicate into two double-stranded helices, with each helix having one of the original DNA strands. This DNA replication was firmly established by 1961 after further demonstration in other species,[208] and of the stepwise chemical reaction.[209][210] According to the 1961 Crick–Monod letter, this experimental proof, along with Wilkins having initiated the DNA diffraction work, were the reasons why Crick felt that Wilkins should be included in the DNA Nobel Prize.[211]

In 1962 the Nobel Prize was subsequently awarded to Crick, Watson, and Wilkins.[9][212][213] Nobel rules prohibit posthumous nominations or splitting of Prizes more than three ways.[214][215] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.[216] By the time of the award Wilkins had been working on the structure of DNA for more than 10 years, and had done much to confirm the Watson–Crick model.[217] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.[218] Watson has suggested that ideally Wilkins and Franklin would have been awarded the Nobel Prize in Chemistry.[7]

Aaron Klug, Franklin's colleague and principal beneficiary in her will, was the sole winner of the Nobel Prize in Chemistry 1982, "for his development of crystallographic electron microscopy and his structural elucidation of biologically important nucleic acid-protein complexes."[219] This work was exactly what Franklin had started and which she introduced to Klug, and it is highly plausible that, were she alive, she would have shared the Nobel Prize.[220]

• 1982, Iota Sigma Pi designated Franklin a National Honorary Member.[221]

Mural inscription on King's College London's Franklin-Wilkins Building, co-named in honour of Rosalind Franklin's work

Blue plaque on 107 Drayton Gardens, London SW10

• 1992, English Heritage placed a blue plaque commemorating Franklin on the building in Drayton Gardens, London, where she lived until her death.[222][223]
• 1993, King's College London renamed the Orchard Residence at their Hampstead Campus as Rosalind Franklin Hall.[224]
• 1993, King's College London placed a blue plaque on its outside wall bearing the inscription: "R. E. Franklin, R. G. Gosling, A. R. Stokes, M. H. F. Wilkins, H. R. Wilson – King's College London – DNA – X-ray diffraction studies – 1953."[225]
• 1995, Newnham College, Cambridge opened a graduate residence named Rosalind Franklin Building,[226] and put a bust of her in its garden.[227][228]
• 1997, Birkbeck, University of London School of Crystallography opened the Rosalind Franklin Laboratory.[229]
• 1997, the asteroid discovered in 1997 was named 9241 Rosfranklin.
• 1998, National Portrait Gallery in London added Rosalind Franklin's portrait next to those of Francis Crick, James Watson and Maurice Wilkins.[230]
• 1999, the Institute of Physics at Portland Place, London, renamed its theatre as Franklin Lecture Theatre.[231]
• 2000, King's College London opened the Franklin–Wilkins Building in honour of Franklin's and Wilkins's work at the college.[232]
• 2001, the American National Cancer Institute established the Rosalind E. Franklin Award for women in cancer research.[233]
• 2002, the University of Groningen, supported by the European Union, launched the Rosalind Franklin Fellowship to encourage women researchers to become full university professors.[234][235]
• 2003, the Royal Society established the Rosalind Franklin Award (officially the Royal Society Rosalind Franklin Award and Lecture) for an outstanding contribution to any area of natural science, engineering or technology.[236] The award consists of a silver-coated medal and a grant of £30,000.[237]
• 2003, the Royal Society of Chemistry declared King's College London as "National Historic Chemical Landmark" and placed a plaque on the wall near the entrance of the building, with the inscription: "Near this site Rosalind Franklin, Maurice Wilkins, Raymond Gosling, Alexander Stokes and Herbert Wilson performed experiments that led to the discovery of the structure of DNA. This work revolutionised our understanding of the chemistry behind life itself."[238]

Rosalind Franklin University of Medicine and Science at Illinois

• 2004, Finch University of Health Sciences/The Chicago Medical School, located in North Chicago, Illinois, USA changed its name to the Rosalind Franklin University of Medicine and Science.[239] It also adopted a new motto "Life in Discovery", and Photo 51 as its logo.[240]
• 2004, the Gruber Foundation started the Rosalind Franklin Young Investigator Award for two female geneticists from all over the world. It carries an annual fund of $25,000, each award is for three years, and selection is made by a joint committee appointed by the Genetics Society of America and the American Society of Human Genetics.[241]
• 2004, the Advanced Photon Source (APS) and the APS Users Organization (APSUO) started the APSUO Rosalind Franklin Young Investigator Award for young scientists who made contributions through the APS.[242]
• 2005, the DNA sculpture (donated by James Watson) outside Clare College, Cambridge's Memorial Court incorporates the words "The double helix model was supported by the work of Rosalind Franklin and Maurice Wilkins."[243]
• 2006, the Rosalind Franklin Society was established in New York by Mary Ann Liebert.[244][245] The Society aims to recognise, foster, and advance the important contributions of women in the life sciences and affiliated disciplines.[246]
• 2008, Columbia University awarded an honorary Louisa Gross Horwitz Prize to Franklin, "for her seminal contributions to the discovery of the structure of DNA".[247]
• 2008, the Institute of Physics established a biennial award the Rosalind Franklin Medal and Prize.[248]
• St Paul's Girls School established the Rosalind Franklin Technology Centre.[25]
• 2012, the bioinformatics education software platform Rosalind is named in honour of Franklin.[249]
• 2012, The Rosalind Franklin Building is opened at Nottingham Trent University.[250]
• 2013, Google honoured Rosalind Franklin with a doodle, showing her gazing at a double helix structure of DNA with an X-ray of Photo 51 beyond it.[251][252]
• 2013, a plaque was placed on the wall of The Eagle pub in Cambridge commemorating Franklin's contribution to the discovery of the structure of DNA, on the sixtieth anniversary of Crick and Watson's announcement in the pub.[253][254]
• 2014, the Rosalind Franklin Award for Leadership in Industrial Biotechnology was established by Biotechnology Industry Organization in collaboration with the Rosalind Franklin Society, for an outstanding woman in the field of industrial biotechnology and bioprocessing.[255]
• 2014, the Rosalind Franklin University of Medicine and Science unveiled a bronze statue of Franklin, created by Julie Rotblatt-Amrany, near its front entrance.[256]
• 2014, the Rosalind Franklin STEM Elementary was opened in Pasco, Washington, the first science, technology, engineering, and math (STEM) elementary school in the district.[257][258]
• 2014, the University of Wolverhampton opened its new laboratory building named the Rosalind Franklin Science Building.[259][260]
• 2015, Newnham College Boat Club, Cambridge, launched a new racing VIII, naming it the Rosalind Franklin.[261]
• 2015, the Rosalind Franklin Appathon was launched by University College London as a national app competition for women in STEMM (science, technology, engineering, maths and medicine).[262]
• 2015, a high performance computing and cloud facility in London was named Rosalind.[263]
• 2016, the British Humanist Association added the Rosalind Franklin Lecture to its annual lecture series, aimed to explore and celebrate the contribution of women towards the promotion and advancement of humanism.[264]
• 2016, the Rosalind Franklin Prize and Tech Day was held on 23 February in London, organised by University College London, i-sense, UCL Enterprise, the London Centre for Nanotechnology and the UCL Athena Swan Charter.[265]
• 2017, DSM opened the Rosalind Franklin Biotechnology Center in Delft, the Netherlands.[266]
• 2017, the Historic England listed the tomb of Franklin under the Planning (Listed Buildings and Conservation Areas) Act 1990 as a "special architectural or historic interest". Official description states that "the tomb commemorates the life and achievements of Rosalind Franklin, a scientist of exceptional distinction, whose pioneering work helped lay the foundations of molecular biology; Franklin’s X-ray observation of DNA contributed to the discovery of its helical structure."[267]
• 2018, the Rosalind Franklin Institute, an autonomous medical research centre under the join venture of 10 universities and funded by the United Kingdom Research and Innovation, was launched at the Harwell Science and Innovation Campus on 6 June.[268]
• 2019, the European Space Agency (ESA) named their ExoMars rover Rosalind Franklin.[269]
• 2019, the University of Portsmouth announced that it changed the name James Watson Halls to Rosalind Franklin Halls from 2 September.[270]

Franklin's part in the discovery of the nature of DNA was shown in the 1987 TV Movie Life Story, starring Juliet Stevenson as Franklin. With Tim Pigott-Smith as Crick, Alan Howard as Wilkins and Jeff Goldblum as Watson. This movie portrayed Franklin as somewhat stern, but also alleged that Watson and Crick did use a lot of her work to do theirs.[271][272]

A 56-minute documentary of the life and scientific contributions of Franklin, DNA – Secret of Photo 51, was broadcast in 2003 on PBS Nova.[273] Narrated by Barbara Flynn, the program features interviews with Wilkins, Gosling, Klug, Maddox,[274] including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley.[275] The UK version produced by BBC is titled Rosalind Franklin: DNA's Dark Lady.[276]

The first episode of another PBS documentary serial, DNA, was aired on 4 January 2004.[277] The episode titled The Secret of Life centres much around the contributions of Franklin. Narrated by Jeff Goldblum, it features Watson, Wilkins, Gosling and Peter Pauling (son of Linus Pauling).[278]

A play titled Rosalind: A Question of Life was written by Deborah Gearing to mark the work of Franklin, and was first performed on 1 November 2005 at the Birmingham Repertory Theatre,[279] and published by Oberon Books in 2006.[280]

Another play, Photograph 51 by Anna Ziegler, published in 2011,[281] has been produced at several places in the US,[282] and in late 2015 was put on at the Noel Coward Theatre, London, with Nicole Kidman playing Franklin.[283] Ziegler's version of the 1951–53 'race' for the structure of DNA sometimes emphasizes the pivotal role of Franklin's research and her personality. Although sometimes altering history for dramatic effect, the play nevertheless illuminates many of the key issues of how science was and is conducted.[284]

False Assumptions by Lawrence Aronovitch is a play about the life of Marie Curie in which Franklin is portrayed as frustrated and angry at the lack of recognition for her scientific contributions.[285]