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Rosalind Elsie Franklin
Rosalind Franklin.jpg
Born Rosalind Elsie Franklin
(1920-07-25)25 July 1920
Notting Hill, London, England
Died 16 April 1958(1958-04-16) (aged 37)
Chelsea, London, England
Ovarian cancer
Resting place Willesden United Synagogue Cemetery
51°32′41″N 0°14′24″W / 51.5447°N 0.2399°W / 51.5447; -0.2399Coordinates: 51°32′41″N 0°14′24″W / 51.5447°N 0.2399°W / 51.5447; -0.2399
Nationality British
Fields Physical chemistry
X-ray crystallography
Institutions British Coal Utilisation Research Association
Laboratoire Central des Services Chimiques de l'État
King's College, London
Birkbeck College, London
Alma mater Newnham College, Cambridge
Thesis The physical chemistry of solid organic colloids with special reference to coal (1945)
Known for Structure of DNA
Fine structure of coal and graphite
Structure of viruses

Rosalind Elsie Franklin (25 July 1920 – 16 April 1958)[1] was an English chemist and X-ray crystallographer who made critical contributions to the understanding of the fine molecular structures of DNA (deoxyribonucleic acid), RNA (ribonucleic acid), viruses, coal, and graphite.[2] Her DNA work achieved the most fame because DNA plays an essential role in cell metabolism and genetics, and the discovery of its structure by Watson and Crick helped them understand how genetic information passes from parents to their offspring.

Franklin is best known for her work on the X-ray diffraction images of DNA, which led to the discovery of the DNA double helix. According to Francis Crick, her data and research were key in determining the structure.[3] Watson confirmed this opinion in his own statement at the opening of the King's College London Franklin–Wilkins building in 2000 and formulating Crick and James Watson's 1953 model regarding the structure of DNA.[4] Franklin's images of X-ray diffraction, confirming the helical structure of DNA, were imprudently shown to Watson by Wilkins. This image provided valuable insight into the DNA structure, but Franklin's scientific contributions to the discovery of the double helix are often overlooked.[5]

Unpublished drafts of her papers (written just as she was arranging to leave King's College, London) show that she had independently determined the overall B-form of the DNA helix and the location of the phosphate groups on the outside of the structure. Moreover, it was a report of Franklin's that convinced Crick and Watson that the backbones had to be on the outside,[5] which was crucial since before this both they and Linus Pauling had independently generated non-illuminating models with the chains inside and the bases pointing outwards.[6] However, her work was published third, in the series of three DNA Nature articles, led by the paper of Watson and Crick which only hinted at her contribution to their hypothesis.[7] Watson has suggested that Franklin would have ideally been awarded a Nobel Prize in Chemistry, along with Maurice Wilkins.[8]

After finishing her portion of the work on DNA, Franklin led pioneering work on the tobacco mosaic virus and the polio virus.[9] She died in 1958 at the age of 37 of ovarian cancer.

Early life and education

Franklin was born in 50 Chepstow Villas,[10] Notting Hill, London into an affluent and influential British Jewish family.[11][12][13] Her father was Ellis Arthur Franklin (1894–1964), a politically liberal London merchant banker who taught at the city's Working Men's College, and her mother was Muriel Frances Waley (1894–1976). Rosalind was the elder daughter and the second child in the family of five children. Her father's uncle was Herbert Samuel (later Viscount Samuel), who was the Home Secretary in 1916 and the first practising Jew to serve in the British Cabinet.[14] Her aunt, Helen Caroline Franklin, was married to Norman de Mattos Bentwich, who was the Attorney General in the British Mandate of Palestine.[15] She was active in trade union organisation and the women's suffrage movement, and was later a member of the London County Council.[16][17] Franklin herself later became an agnostic.[18][19] Her uncle, Hugh Franklin, was another prominent figure in the suffrage movement, although his actions therein embarrassed the Franklin family.[13]

From early childhood, Franklin showed exceptional scholastic abilities. She was educated at St Paul's Girls' School[20][21] where she excelled in science, Latin,[22] and sports.[23] Her family was actively involved with in the Working Men's College, where her father taught the subjects of electricity, magnetism, and the history of the Great War in the evenings, later becoming the vice-principal.[24][25] Later Franklin's family helped settle Jewish refugees from Europe who had escaped the Nazis.[17]

Cambridge, Kingston and Paris

Franklin went to Newnham College, Cambridge in 1938 and studied chemistry within the Natural Sciences Tripos. Her teachers included the spectroscopist W.C. Price, who later became one of her senior colleagues at King's College.[26] 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.[27]

Franklin was there awarded a research fellowship. According to an entry on the web site of the Dolan DNA Learning Center of the Cold Spring Harbor Laboratory, "She spent a year in R.G.W. Norrish's lab without great success."[28] Resigning from Norrish's Lab, Franklin fulfilled the requirements of the National Service Act by working as an Assistant Research Officer at the British Coal Utilisation Research Association (BCURA).[9] The BCURA was located on the Coombe Springs Estate near Kingston upon Thames near the southwestern boundary of London. Professor 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 attended.[2]

She studied the porosity of coal and compared helium's density. 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, Franklin helped classify coals and accurately predict their performance for fuel purposes and for production of wartime devices (i.e. gas masks).[29] This work was the basis of her Ph.D. thesis The physical chemistry of solid organic colloids with special reference to coal for which Cambridge University awarded her a Ph.D. in 1945. It was also the basis of several papers.[2]

The French scientist Adrienne Weill was one of Franklin's tutors at Newnham. At the end of World War II, according to Anne Sayre, author of Rosalind Franklin and DNA, Franklin asked Weill 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 Franklin 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 Franklin's appointment with Jacques Mering at the Laboratoire Central des Services Chimiques de l'Etat in Paris. Franklin joined the labo (as referred to by the staff) of Mering on 14 February 1947 as one of the fifteen chercheurs (researchers).[30][31]

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, in particular the changes to the arrangement of atoms when it is converted to graphite.[2] Franklin published several further papers on this work. It became part of the mainstream of work on the physics and chemistry of coal, covered by a current monograph,[32] the annual[33] and other publications. Mering also continued the study of carbon in various forms, using X-ray diffraction and other methods.[34]

King's College, London

In January 1951, Franklin started working as a research associate at King's College, London in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall.[35] Although originally she was to have worked on X-ray diffraction of proteins and lipids in solution, Randall redirected her work to DNA fibres before she started working at King's since Franklin was to be the only experienced experimental diffraction researcher at King's in 1951.[36][37] He made this reassignment, even before she started working at King's, because of the following pioneering work by Maurice Wilkins and Raymond Gosling – a Ph.D. student assigned to help Franklin.[38][39]
Even using crude equipment, these two men had obtained an outstanding diffraction picture of DNA which sparked further interest in this molecule. Wilkins and Gosling 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.[40]
Randall's lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.[41]

Franklin, working with Gosling,[42] 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, Franklin also skillfully manipulated the critical hydration of her specimens.[43] When Wilkins inquired about this improved technique, Franklin replied in terms which offended Wilkins as Franklin had "an air of cool superiority".[44]

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.[45] 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.[46][47]

These forms were termed DNA "B" and "A" respectively. Because of the intense personality conflict developing between Franklin and Wilkins, Randall[48] divided the work on DNA. Franklin chose the data rich A form while Wilkins selected the "B" form[49][50] because his preliminary pictures had hinted it might be helical. He showed tremendous insight in this assessment of preliminary data. The X-ray diffraction pictures taken by Franklin at this time have been called, by J. D. Bernal, as "amongst the most beautiful X-ray photographs of any substance ever taken".[46]

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 1952 May, Franklin became unconvinced that the A form of DNA was helical in structure.[51] 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 death notice regretting the 'death' of helical crystalline DNA (A-DNA).[52] During 1952, Rosalind Franklin and Raymond Gosling worked at applying the Patterson function to the X-ray pictures of DNA they had produced.[53] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.[54][55]

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 form manuscripts reached Acta Crystallographica in Copenhagen on 6 March 1953,[56] one day before Crick and Watson had completed their model.[57] Franklin must have mailed them while the Cambridge team was building their model, and certainly had written them before she knew of their work. 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.[58]

The third draft paper on the "B" form of DNA, dated 17 March 1953, was discovered years later amongst her papers, 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.[59] Klug designed this paper to complement the first article he had written defending Franklin's significant contribution to DNA structure.[60] 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 changed the course of DNA history with the following disclosure. Wilkins imprudently showed Watson Franklin's DNA X-ray image. Watson, in turn, showed Wilkins a prepublication manuscript by Pauling and Corey.[61] Franklin and Gosling's photo 51 gave the Cambridge pair critical insights into the DNA structure, whereas Pauling and Corey's paper described a molecule remarkably like their first incorrect model.

DNA modelling

In February 1953, Francis Crick and James D. Watson of the Cavendish Laboratory in Cambridge University had started to build a model of the B form of DNA using data similar to that available to both teams at King's. Much of their data were 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 were critical.[62] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,[49][63] but Franklin was opposed to prematurely building theoretical models, until sufficient data were obtained properly to guide the model building. She took the view that building a model was to be undertaken only after enough of the structure was known.[51][64]

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.[65]

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".[66] However they knew they must complete their model before they could be certain.[67]

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".[68] This was also one day after Franklin's two A form 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.[69]

It is uncertain how long it took for Gosling to inform Franklin at Birkbeck, but her original 17 March B form 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,[70] 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".[71]

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 Wilkin's 'unpublished' contribution.[4] 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.[72][73]

Weeks later, on 10 April, Franklin wrote to Crick for permission to see their model.[74] 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.[75]

However, as documented above, she did not hesitate to publish preliminary ideas about DNA in Acta, even before they could be definitively proven. 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.[76][77][78]

Birkbeck College


An electronmicrograph 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, to which she described (in a letter to Adriene Weill in Paris) as "moving from a palace to the slums... but pleasanter all the same."[79] She was recruited by physics department chair J. D. Bernal,[80] a brilliant crystallographer who happened to be an Irish communist, known for promoting women crystallographers. She worked as a senior scientist with her own research group, funded by the Agricultural Research Council (ARC).[81] Despite the parting words of Bernall 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 25 July issue of Nature.[82] Moreover, she continued to venture into 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 the early 1954 proved to be a longstanding and successful collaboration. Klug had just then earned his PhD from Trinity College, Cambridge, and joined Birkbeck in the late 1953. In 1955 Franklin published her first major works on TMV in Nature, in which she described that TMV virus particles were all of the same length.[83] This was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct.[84]

Franklin assigned the study of the complete structure of TMV to her PhD student Kenneth Holmes. They soon discovered (published in 1956) that the covering of TMV was protein molecules arranged in helices.[85] Her colleague Klug worked on spherical viruses with his student John Finch, with Franklin coordinating and overseeing the work.[86] As a team, from 1956 they started publishing seminal works on TMV,[87] cucumber virus 4 and turnip yellow mosaic virus.[88]

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.[89] The Birkbeck team members continued working on RNA viruses affecting several plants, including potato, turnip, tomato and pea.[90] 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 complimentary papers in 10 March issue of Nature, in which they showed that the RNA in TMV is wound along the inner surface of the hollow virus.[91][92] Casper was not an enthusiastic writer such that Franklin had to write the entire manuscript for him.[93]

The first major international fair after World War II, named Expo 58, was to be held in 1958 in Brussels.[94] Franklin was invited to make a five-foot high model of TMV, which she started in 1957. Her materials were ping pong balls and plastic bicycle handlebar grips.[95] The Brussels world's fair, with an exhibit of her virus model at the International Science Pavilion, opened on 17 April, just the day after she died.[96]

Her main research team at Birkbeck College, London Klug, Finch and Holmes moved to the Laboratory of Molecular Biology, Cambridge in 1962.

Illness and death

In mid-1956, while on a work-related trip to the United States, Franklin first began to suspect a health problem.[97] An operation in September of the same year revealed two tumours in her abdomen.[98]
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,[99] 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.[100] In 1957, the group was also working on the polio virus and had obtained funding from the Public Health Service of the National Institutes of Health in the United States for this.[101]

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.[102] She returned to work in January 1958, and she was given a promotion to Research Associate in Biophysics on 25 February.[103] She fell ill again on 30 March, and she died on 16 April 1958, in Chelsea, London,[104][105] of bronchopneumonia, secondary carcinomatosis, and ovarian cancer. Exposure to X-ray radiation is sometimes considered to be a possible factor in her illness.[106]

Other members of her family have died of cancer, and the incidence of gynaecological cancer is known to be disproportionately high among Ashkenazi Jews.[107] Her death certificate read: A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.[108] She was interred on 17 April 1958 in the family plot at Willesden United Synagogue Cemetery at Beaconsfield Road in London Borough of Brent.[109] The inscription on her tombstone reads:[110][111]
IN MEMORY OF
ROSALIND ELSIE FRANKLIN
מ' רחל בת ר' יהודה
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"]

Controversies after death

Various controversies surrounding Rosalind Franklin came to light following her death.

Allegations of sexism

Sayre, one of Franklin's biographers, states "In 1951 ... King's College 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" (page 96, lines 8 to 10; page 97, line 4;page 99, footnote). Andrzej Stasiak states "Sayre's book became widely cited in feminist circles for exposing rampant sexism in science."[112] 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".[113] Maddox states, (page 134, final paragraph) "Randall ... did have many women on his staff ... they found him ... sympathetic and helpful."

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".[114][115] Elkin states that most of the MRC group typically ate lunch together (including Franklin) in the mixed dining room discussed below.[116] 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."

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.[117] In this, he denigrates her work and frequently refers to her in patronizing terms as "Rosy", a name she never used. Much later, at Cambridge, Francis Crick acknowledges, "I'm afraid we always used to adopt – let's say, a patronizing attitude towards her." And another Cambridge colleague, Peter Cavendish, wrote in a letter, "Wilkins is supposed to be doing this work; Miss Franklin is evidently a fool."

Franklin herself is said to have been "not immune to the sexism rampant in these circles." In a letter to her parents in January 1939, she called one lecturer "very good, though female."[118][dubious – discuss]

Contribution to the model of DNA

Rosalind Franklin's first important contributions to the Crick and Watson model 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, and her position whereby 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.[119]

Photo 51 taken by Raymond Gosling in May 1952.

The other contribution included an X-ray photograph of B-DNA (called photograph 51),[120] that was briefly shown to Watson by Wilkins in January 1953,[121][122] 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.[123][124]

Started by Sayre,[125] a story has been reiterated that the photograph 51 in question was shown to Watson by Wilkins – without Franklin's permission,[112][126][127] and that it was a case of bad science ethics.[128] But it is not true. Wilkins had been given photograph 51 by Franklin's Ph.D. student Gosling, because she was leaving King's to work at Birkbeck. There was allegedly nothing untoward in this transfer of data to Wilkins,[129][130] since the 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.[131] Also it was implied by Horace Freeland Judson, incorrectly, that Maurice Wilkins had taken the photograph out of Franklin's drawer.[132]

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".[133] 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[134] and to post a general statement in Science excusing himself on the basis of being "inexperienced and casual in administrative matters".[135]

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.[136][137]

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. For a long time it was generally believed that Franklin was never aware that her work had been used during construction of the model,[138] but Gosling revealed in his 2013 interview that, "Yes. Oh, she did know about that."[139]

Recognition of her contribution to the model of DNA

Upon the completion of their model, Crick and Watson had invited Wilkins to be a co-author of their paper describing the structure.[140][79] Wilkins turned down this offer, as he had taken no part in building the model.[141] 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.[142] There is no doubt that Franklin's experimental data were used by Crick and Watson to build their model of DNA in 1953 (see above). Some, including Maddox as cited next, 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.[143]

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".[144]

Twenty five 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.[145] 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.[146]

Sayre's early analysis was often ignored because of perceived feminist overtones in her book. It should be noted that in their original paper, Watson and Crick do not cite the X-ray diffraction work of both Wilkins and Franklin. However, they admit their 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."[4] Watson and Crick had no experimental data to support their model. It was Franklin and Gosling's own publication in the same issue of Nature with the X-ray image of DNA, which served as the main evidence; in which they concluded:
Thus our general ideas are not inconsistent with the model proposed by Watson and Crick in the preceeding communication.[147]

Nobel Prize

Franklin was never nominated for a Nobel Prize.[148][149] She had died in 1958, and during her lifetime the DNA structure was not considered as 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.[150] The first major breakthrough was from Matthew Meselson and Franklin Stahl in 1958, who experimentally showed the DNA replication of a bacterium Escherichia coli.[151] 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,[152] and of the stepwise chemical reaction.[153][154] 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.[3]

In 1962 the Nobel Prize was subsequently awarded to Crick, Watson, and Wilkins.[9][155][156] It is not clear whether she would have been included, had she lived.[157] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.[158] 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.[159] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.[160] Watson has suggested that ideally Wilkins and Franklin would have been awarded the Nobel Prize in Chemistry.[8]

It is also interesting to note that Franklin's colleague and principal beneficiary in her will, Klug 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."[161] This work was exactly what Franklin had started and which she introduced it to Klug, and it is highly plausible that, were she alive, she would have shared the Nobel prize.[162]

Posthumous recognition

  • 1982, Iota Sigma Pi designated Franklin a National Honorary Member.[163]

    Blue plaque on SW10, Drayton Gardens, Donovan Court
  • 1992, English Heritage placed a blue plaque on the entrance wall of SW10, Drayton Gardens, 107, Donovan Court, where Rosalind Franklin lived during her professional career till her death.[164] The inscription reads "Rosalind Franklin, 1920-1958, pioneer of the study of molecular structures including DNA, lived here, 1951-1958".[165]
  • 1993, King's College London renamed the Orchard Residence at their Hampstead Campus on Kidderpore Avenue as Rosalind Franklin Hall.[166]
  • 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."[167]
  • 1995, Newnham College opened a graduate residence named Rosalind Franklin Building[168] and put a bust of her in its garden.[169][170]
  • 1997, Birkbeck, University of London School of Crystallography opened the Rosalind Franklin Laboratory.[171]
  • 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.[172]
  • 1999, the Institute of Physics at Portland Place, London, renamed its theatre as Franklin Lecture Theatre.[173]
  • 2000, King's College London opened the Franklin–Wilkins Building in honour of Dr. Franklin's and Professor Wilkins's work at the college.[174] King's had earlier, in 1994, also named one of the Halls in Hampstead Campus residences in memory of Rosalind Franklin.
  • 2001, the American National Cancer Institute established the Rosalind E. Franklin Award for Women in Science.[175]
  • 2003, the Royal Society established the Rosalind Franklin Award, for an outstanding contribution to any area of natural science, engineering or technology.[176]
  • 2003, the Royal Society of Chemistry declared King's College London as "National Historic Chemical Landmark" and placed a plague on the wall near the entrance of the building, with an epitaph: "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."[177]

    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.[178]
  • 2005, the wording on the DNA sculpture (which was donated by James Watson) outside Clare College, Cambridge's Memorial Court is a) on the base: i) "These strands unravel during cell reproduction. Genes are encoded in the sequence of bases." and ii) "The double helix model was supported by the work of Rosalind Franklin and Maurice Wilkins.", as well as b) on the helices: i) "The structure of DNA was discovered in 1953 by Francis Crick and James Watson while Watson lived here at Clare." and ii) "The molecule of DNA has two helical strands that are linked by base pairs Adenine – Thymine or Guanine – Cytosine."[179]
  • 2006, the Rosalind Franklin Society was established in New York,[180] which aims to recognise, foster, and advance the important contributions of women in the life sciences and affiliated disciplines.[181]
  • 2007, the University of Groningen, supported by the European Union, launched the Rosalind Franklin Fellowship to encourage women researchers to become full university professors.[182][183]
  • 2008, Columbia University awarded an Honorary Louisa Gross Horwitz Prize to Rosalind Franklin, Ph.D., posthumously, "for her seminal contributions to the discovery of the structure of DNA".[184]
  • 2012, honoured as the namesake of Rosalind, an online project teaching programming via molecular biology.
  • 2012, The Right Honourable Professor The Lord Robert Winston opens the Nottingham Trent University multi-million pound research building the Rosalind Franklin Building.[185]
  • 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.[186][187]
  • 7 March 2013, a plaque was placed on the wall of The Eagle, a pub near Cambridge University where Crick and Watson announced the discovery of the structure of DNA sixty years to the day.[188][189]
  • 2014, BIO Rosalind Franklin Award 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.[190]

In popular culture

Franklin's part in the discovery of the nature of DNA was shown in the 1987 TV Movie Life Story, starring Tim Pigott-Smith as Crick, Alan Howard as Wilkins, Jeff Goldblum as Watson, and Juliet Stevenson as Franklin. This movie portrayed Franklin as somewhat stern, but also alleged that Watson and Crick did use a lot of her work to do theirs.[191][192]

A 56-minute documentary of the life and scientific contributions of Franklin, DNA - Secret of Photo 51, was broadcast in 2003 on PBS NOVA.[193] The program features interviews with Wilkins, Gosling, Klug, Maddox,[194] including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley.[195]

The first episode of another PBS documentary serial, DNA, was aired on 4 January 2004.[196] 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).[197]

A play entitled 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,[198] and published by Oberon Books in 2006.[199]

Another play, Photograph 51 by Anna Ziegler, published in 2011,[200] has been produced at several places in the USA.[201] 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.[202]

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.[203]

Publications

Rosalind Franklin produced a number of publications, some cited a number of times. A representative sample is listed below. The last two publications in this list were published posthumously.