Rosalind Franklin

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Rosalind Elsie Franklin
Rosalind Franklin.jpg
BornRosalind Elsie Franklin
(1920-07-25)25 July 1920
Notting Hill, London, England
Died16 April 1958(1958-04-16) (aged 37)
Chelsea, London, England
Ovarian cancer
Resting placeWillesden 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
FieldsPhysical chemistry
X-ray crystallography
InstitutionsBritish Coal Utilisation Research Association
Laboratoire Central des Services Chimiques de l'État
King's College, London
Birkbeck College, London
Alma materNewnham College, Cambridge
ThesisThe physical chemistry of solid organic colloids with special reference to coal (1945)
Known forStructure of DNA
Fine structure of coal and graphite
Structure of viruses
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Rosalind Elsie Franklin
Rosalind Franklin.jpg
BornRosalind Elsie Franklin
(1920-07-25)25 July 1920
Notting Hill, London, England
Died16 April 1958(1958-04-16) (aged 37)
Chelsea, London, England
Ovarian cancer
Resting placeWillesden 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
FieldsPhysical chemistry
X-ray crystallography
InstitutionsBritish Coal Utilisation Research Association
Laboratoire Central des Services Chimiques de l'État
King's College, London
Birkbeck College, London
Alma materNewnham College, Cambridge
ThesisThe physical chemistry of solid organic colloids with special reference to coal (1945)
Known forStructure 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[edit]

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[edit]

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[edit]

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[edit]

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[edit]

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[edit]

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]

מ' רחל בת ר' יהודה
25TH JULY 1920 - 16TH APRIL 1958
ת נ צ ב ה [Hebrew initials for "her soul shall be bound in the bundle of life"]

Controversies after death[edit]

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

Allegations of sexism[edit]

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 ]

Contribution to the model of DNA[edit]

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[edit]

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[edit]

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[edit]

In popular culture[edit]

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] Narrated by Barbara Flynn, the program features interviews with Wilkins, Gosling, Klug, Maddox,[194] including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley.[195] The UK version produced by BBC is titled Rosalind Franklin: DNA's Dark Lady.[196]

The first episode of another PBS documentary serial, DNA, was aired on 4 January 2004.[197] 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).[198]

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,[199] and published by Oberon Books in 2006.[200]

Another play, Photograph 51 by Anna Ziegler, published in 2011,[201] has been produced at several places in the USA.[202] 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.[203]

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.[204]


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.


  1. ^ "The Rosalind Franklin Papers, Biographical Information". Retrieved 13 November 2011. 
  2. ^ a b c d e "The Rosalind Franklin Papers, The Holes in Coal: Research at BCURA and in Paris, 1942–1951". Retrieved 13 November 2011. 
  3. ^ a b Zallen, Doris T. (2003). "Despite Franklin's work, Wilkins earned his Nobel". Nature 425 (6953): 15. Bibcode:2003Natur.425...15Z. doi:10.1038/425015b. PMID 12955113. (Crick's 31 December 1961 letter to Jacques Monod) However, the data which really helped us to obtain the structure was mainly obtained by Rosalind Franklin 
  4. ^ a b c Watson JD, Crick FH (April 1953). "Molecular structure of nucleic acids; a structure for deoxyribose nucleic acid" (PDF). Nature 171 (4356): 737–738. Bibcode:1953Natur.171..737W. doi:10.1038/171737a0. PMID 13054692.  This article was immediately followed by the two King's submissions: Wilkins MHF, Stokes AR, Wilson HR (April 1953). "Molecular structure of deoxypentose nucleic acids". Nature 171 (4356): 738–740. Bibcode:1953Natur.171..738W. doi:10.1038/171738a0. PMID 13054693.  then by: Franklin RE, Gosling RG (April 1953). "Molecular configuration in sodium thymonucleate" (PDF). Nature 171 (4356): 740–741. Bibcode:1953Natur.171..740F. doi:10.1038/171740a0. PMID 13054694. 
  5. ^ a b [1] Rosalind Franklin's Legacy, Interview of Lynn Osman Elkin conducted on 26 March 2003
  6. ^ In Pursuit of the Gene. From Darwin to DNA — By James Schwartz. Harvard University Press, 2008
  7. ^ "''Double Helix: 50 Years of DNA.'' Nature archives.". Nature Publishing Group. Retrieved 2013-07-25. 
  8. ^ a b "The Discovery of the Molecular Structure of DNA - The Double Helix". Official Website of the Nobel Prizes. Retrieved 4 February 2014. 
  9. ^ a b c "James Watson, Francis Crick, Maurice Wilkins, and Rosalind Franklin". Chemical Heritage Foundation. Retrieved 1 November 2013. 
  10. ^ "Name of Firm: A. Kyeser & Co.". The Gazette. 22 February 1922. Retrieved 21 November 2014. 
  11. ^ "Rosalind Franklin". London Remembers. Retrieved 21 November 2014. 
  12. ^ GRO Register of Births: SEP 1920 1a 250 KENSINGTON – Rosalind E. Franklin, mmn = Waley
  13. ^ a b Maddox, Brenda (2002). Rosalind Franklin: The Dark Lady of DNA. HarperCollins. ISBN 0-06-018407-8. 
  14. ^ Maddox p. 7
  15. ^ Segev p.
  16. ^ Sayre, A. (1975). Rosalind Franklin and DNA. New York: Norton. p. 31. ISBN 0393074935. 
  17. ^ a b Maddox p. 40
  18. ^ "This flat declaration prompted Ellis Franklin to accuse his strong-willed daughter of making science her religion. He was right. Rosalind sent him a four-page declaration, eloquent for a young woman just over 20, let alone a scientist of any age. ..."It has just occurred to me that you may raise the question of a creator. A creator of what? […] I see no reason to believe that a creator of protoplasm or primeval matter, if such there be, has any reason to be interested in our insignificant race in a tiny corner of the universe, and still less in us, as still more insignificant individuals. Again, I see no reason why the belief that we are insignificant or fortuitous should lessen our faith – as I have defined it."" Brenda Maddox, Mother of DNA, – Volume 117 Issue 3 Autumn 2002.
  19. ^ Listed as an agnostic on Rosalind Franklin,
  20. ^ Maddox p. 25
  21. ^ Sayre p. 41
  22. ^ Maddox p. 30
  23. ^ Maddox, p. 26
  24. ^ Maddox, p. 20
  25. ^ Sayre, p. 35
  26. ^ R.N. Dixon, D.M. Agar and R.E. Burge, William Charles Price. 1 April 1909 – 10 March 1993, Biographical Memoirs of Fellows of the Royal Society, vol. 43, page 438, line 17, 1997.
  27. ^ Fact sheet: Women at Cambridge: A Chronology, [2][dead link].
  28. ^ Rosalind Franklin, Cold Spring Harbor Laboratory's Dolan DNA Learning Center, ID 1649, [3].
  29. ^ "The Rosalind Franklin Papers: The Holes in Coal: Research at BCURA and in Paris, 1942–1951". Retrieved 2013-07-25. 
  30. ^ "Rosalind Franklin". Timetoast. Retrieved 28 August 2014. 
  31. ^ "Rosalind Franklin (1920-1958)". DNA Learning Center, Cold Spring Harbor Laboratory. Retrieved 28 August 2014. 
  32. ^ D. W. van Krevelen, Coal, Third Edition: Typology – Physics – Chemistry – Constitution, Elsevier, New York, 1993.
  33. ^ Chemistry and Physics of Carbon, vol 1–, 1968–, Elsevier, New York.
  34. ^ G. Terriere, A. Oberlin, J. Mering, Oxidation of graphite in liquid medium -- observations by means of microscopy and electron diffraction, Carbon, 5, 431--, 1967.
  35. ^ Maddox, p. 124
  36. ^ Maddox, p. 114
  37. ^ Wilkins, Wilkins, M., The Third Man of the Double Helix, an autobiography (2003) Oxford University Press, Oxford. pp. 143–144
  38. ^ The Dark Lady Of DNA by Branda Maddox
  39. ^ Wilkins, p. 121
  40. ^ Maddox, pp. 149–150, Elkin, p 45. Elkin, L.O. Rosalind Franklin and the Double Helix. Physics Today, March 2003(available free on-line, see references). Olby, R. The Path to the Double Helix (London: MacMillan, 1974).
  41. ^ Sayre, Olby, Maddox, Elkin, Wilkins
  42. ^ Maddox, p. 129
  43. ^ Elkin, p. 43
  44. ^ Wilkins p. 155
  45. ^ Elkin p. 45
  46. ^ a b Maddox, p. 153
  47. ^ Wilkins, p. 154
  48. ^ Maddox p 155
  49. ^ a b Wilkins, p. 158
  50. ^ Maddox, p. 155
  51. ^ a b Wilkins, p. 176
  52. ^ Wilkins, p. 182
  53. ^ Maddox, p. 168
  54. ^ Maddox, p. 169
  55. ^ Wilkins, pp. 232–233
  56. ^ Franklin, R.E. and Gosling, R.G. authors of papers received 6 March 1953 Acta Cryst. (1953). 6, 673 The Structure of Sodium Thymonucleate Fibres I. The Influence of Water Content Acta Cryst. (1953). 6, 678 The Structure of Sodium Thymonucleate Fibres II. The Cylindrically Symmetrical Patterson Function
  57. ^ Maddox p 205
  58. ^ Acta Cryst. (1953). 6, 673 The Structure of Sodium Thymonucleate Fibres I. The Influence of Water Content
  59. ^ Klug, A. "Rosalind Franklin and the Double Helix", Nature 248 (26 April 1974): 787–788
  60. ^ Klug, A. Rosalind Franklin and the Discovery of the Structure of DNA, Nature 219 (24 August 1968): 808–810 & 843.
  61. ^ Yockey, pp. 9–10
  62. ^ Crick's 31 December 1961 letter to Jacque Monod cited above
  63. ^ Maddox, p. 147
  64. ^ Maddox, p. 161
  65. ^ Hubbard, Ruth (1990). The Politics of Women's Biology. Rutgers State University. p. 60. ISBN 0-8135-1490-8. 
  66. ^ "The Double Helix" p. 115
  67. ^ "The Double Helix" p. 60
  68. ^ "All hands to the pump" letter is preserved in the Crick archives at the University of California, San Diego, and was posted as part of their Web collection. It is also quoted by both Maddox, p 204, and Olby.
  69. ^ Maddox p. 207
  70. ^ In contrast to his other letters to Crick, Wilkins dated this one.
  71. ^ "Old rogues" letter is preserved in the Crick archives at the University of California at San Diego, and was posted as part of their Web collection. It is also quoted by both Maddox, p. 208 and Olby.
  72. ^ Franklin and Gosling (1953)
  73. ^ Maddox, p. 210
  74. ^ 10 April 1953 Franklin post card to Crick asking permission to view model. The original is in the Crick archives at the University of California, San Diego.
  75. ^ Holt, J. (2002)
  76. ^ Rich, Alexander (2003). "The double helix: a tale of two puckers". Nature Structural Biology 10 (4): 247–249. doi:10.1038/nsb0403-247. PMID 12660721. 
  77. ^ Scher, Stanley (2004). "Was Watson and Crick's model truly self-evident?". Nature 427 (6975): 584–584. doi:10.1038/427584c. PMID 14961092. 
  78. ^ Arnott, Struther (2006). "Historical article: DNA polymorphism and the early history of the double helix". Trends in Biochemical Sciences 31 (6): 349–354. doi:10.1016/j.tibs.2006.04.004. PMID 16678428. 
  79. ^ a b Maddox, p. 205
  80. ^ Maddox, p. 229
  81. ^ Maddox, p. 235
  82. ^ Franklin, RE; Gosling, RG (1953). "Evidence for 2-chain helix in crystalline structure of sodium deoxyribonucleate.". Nature 172 (4369): 156–157. doi:10.1038/172156a0. PMID 13072614. 
  83. ^ Franklin, RE. (1955). "Structure of Tobacco Mosaic Virus". Nature 175 (4452): 379–381. doi:10.1038/175379a0. PMID 14356181. 
  84. ^ Maddox, p. 252
  85. ^ Franklin and Holmes, 1956
  86. ^ Maddox, p. 254
  87. ^ Franklin, Rosalind E.; Klug, A. (1956). "The nature of the helical groove on the tobacco mosaic virus particle X-ray diffraction studies". Biochimica et Biophysica Acta 19: 403–416. doi:10.1016/0006-3002(56)90463-2. PMID 13315300. 
  88. ^ Franklin et al., 1958
  89. ^ Maddox, p. 256
  90. ^ Maddox, p. 262
  91. ^ Franklin, RE (1956). "Structure of Tobacco Mosaic Virus: Location of the Ribonucleic Acid in the Tobacco Mosaic Virus Particle". Nature 177 (4516): 928–930. doi:10.1038/177928b0. 
  92. ^ Casper, D. L. D. (1956). "Structure of Tobacco Mosaic Virus: Radial Density Distribution in the Tobacco Mosaic Virus Particle". Nature 177 (4516): 928–928. doi:10.1038/177928a0. 
  93. ^ Maddox, p. 269
  94. ^ "Expo 58". Retrieved 21 January 2015. 
  95. ^ "Behind the picture: Rosalind Franklin and the polio model". Medical Research Council. Retrieved 21 January 2015. 
  96. ^ Maddox, Brenda. "Mother of DNA". New Humanist. Retrieved 21 January 2015. 
  97. ^ Maddox, p. 284
  98. ^ Maddox, p. 285
  99. ^ Maddox, p. 288
  100. ^ Maddox, p. 292
  101. ^ Maddox, p. 296
  102. ^ Maddox, p. 301
  103. ^ Maddox, p. 302
  104. ^ GRO Register of Deaths: JUN 1958 5c 257 CHELSEA – Rosalind E. Franklin, aged 37
  105. ^ Maddox, pp. 305–307
  106. ^ "Defending Franklin's Legacy". Secret of Photo 51. NOVA. Retrieved 10 November 2010. Along with genetic predisposition; opinion of CSU's Lynne Osman Elkin; see also March 2003 Physics Today
  107. ^ Maddox, p.320
  108. ^ Murray, Ruby J (2011). "Historical Profile: Rosalind Franklin". Dumbo Feather. Retrieved 27 August 2014. 
  109. ^ "Rosalind Elsie Franklin". Find A Grave. 19 October 2001. Retrieved 27 August 2014. 
  110. ^ "Rosalind Franklin tomb". Himetop. Retrieved 27 August 2014. 
  111. ^ Friedman, Meyer; Friedland, Gerald W. Medicine'S 10 Greatest Discoveries. Universities Press. p. 227. ISBN 9788173712265. 
  112. ^ a b Stasiak, Andrzej (March 2001). "Rosalind Franklin". EMBO Reports 2 (3): 181. doi:10.1093/embo-reports/kve037. PMC 1083834. 
  113. ^ Hussain, Farooq (20 November 1975). Did Rosalind Franklin deserve DNA Nobel prize?. New Scientist 68 (976) (Reed Business Information). p. 470. Retrieved 10 January 2011. 
  114. ^ Sayre, p.97
  115. ^ Bryson, B. (2004) p. 490
  116. ^ Elkin, p.45
  117. ^ Harding, Sandra (2006). "Sexist criticism of Watson's memoir". Science and Social Inequality: Feminist and Postcolonial Issues. Urbana: University of Illinois Press. p. 71. ISBN 978-0-252-07304-5. Retrieved 10 January 2011. 
  118. ^ Wertheimer, Michael; Clamar, Aphrodite; Siderits, Mary Anne (2007). "The Case of the Purloined Picture: Rosalind Franklin and the Keystone of the Double Helix". In Gavin, Eileen A.; Clamar, Aphrodite; Siderits, Mary Anne. Women of Vision: Their Psychology, Circumstances, and Successes. New York: Springer Science+Business Media. ISBN 978-0-8261-0253-9. Retrieved 10 January 2011  Rosalind's letter quoted 
  119. ^ Everson, Ted (2007). The Gene: A Historical Perspective. Westport, Connecticut: Greenwood Press. p. 85. ISBN 978-0-313-33449-8. 
  120. ^ Maddox, pp. 177–178
  121. ^ Maddox, p. 196
  122. ^ Crick, (1988) p. 67.
  123. ^ Elkin, L.O. (2003)p 44
  124. ^ Maddox, pp. 198–199
  125. ^ Sayre, p. 151
  126. ^ Minkoff, Eli; Baker, Pamela (2000). Biology Today: An Issues Approach (2 ed.). New York: Garland Publishing. p. 58. ISBN 978-0-8153-2760-8. 
  127. ^ Creager, Angela (2003). "Crystallizing a Life in Science". American Scientist. Sigma Xi, The Scientific Research Society. Retrieved 25 January 2015. 
  128. ^ Stasiak, Andrzej (2003). "The First Lady of DNA". EMBO Reports 4 (1): 14. doi:10.1038/sj.embor.embor723. PMC 1315822. 
  129. ^ Maddox, pp. 196
  130. ^ Wilkins, p. 198
  131. ^ Maddox p.312,
  132. ^ Wilkins, p. 257
  133. ^ Maddox p.188
  134. ^ Perutz's papers are in the Archive of the J. Craig Venter institute and Science Foundation in Rockville Maryland, which were purchased as part of the Jeremy Norman Archive of Molecular Biology; quoted in Ferry, Georgina, 2007. Max Perutz and the Secret of Life. Published in the UK by Chatto & Windus (ISBN 0-701-17695-4), and in the USA by the Cold Spring Harbor Laboratory Press.
  135. ^ Science, 27 June 1969, pp. 207–212, also reprinted in the Norton critical edition of The Double Helix, edited by Gunther Stent.
  136. ^ Maddox, p. 199
  137. ^ Watson (1969).
  138. ^ Maddox, p. 316
  139. ^ Attar, N (2013). "Raymond Goslin: the man who crystallized genes". Genome Biology 14 (4): 402. doi:10.1186/gb-2013-14-1-402. PMC 3663117. PMID 23651528. 
  140. ^ Wilkins, p. 213
  141. ^ Wilkins, p. 214
  142. ^ Wilkins, p. 226
  143. ^ Maddox, p. 207
  144. ^ Maddox, pp316–317, and other parts of the epilogue
  145. ^ Watson, J.D. (1968) pp. 95–96
  146. ^ Sayre,A. (1975) pp. 156–167
  147. ^ Franklin, Rosalind E.; Gosling, R. G. (April 25, 1953). "Molecular Configuration in Sodium Thymonucleate" (PDF). Nature 171 (4356): 740–741. Bibcode:1953Natur.171..740F. doi:10.1038/171740a0. PMID 13054694. 
  148. ^ "The Discovery of the Molecular Structure of DNA – The Double Helix". 2003-09-30. Retrieved 2013-07-25. 
  149. ^ Washington, Harriet A. (2012-12-31). "Don't Forget Rosalind Franklin". Ms. 
  150. ^ Gann, Alexander; Witkowski, Jan A. (2013). "DNA: Archives reveal Nobel nominations". Nature 496 (7446): 434. doi:10.1038/496434a. PMID 23619686. 
  151. ^ Meselson, Matthew; Stahl, Franklin W. (1958). "The replication of DNA in Escherichia coli". Proceedings of the National Academy of Sciences 44 (7): 671–682. doi:10.1073/pnas.44.7.671. PMC 528642. PMID 16590258. 
  152. ^ Nakada, D; Ryan, FJ (1961). "Replication of deoxyribonucleic acid in non-dividing bacteria". Nature 189: 398–399. doi:10.1038/189398a0. PMID 13727575. 
  153. ^ Dounce, AL; Sarkar, NK; Kay, ER (1961). "The possible role of DNA-ase I in DNA replication". Journal of Cellular and Comparative Physiology 57 (1): 47–54. PMID 13724093. 
  154. ^ Cavalleiri, LF; Rosenberg, BH (1961). "The replication of DNA III. Changes in the number of strands in E. coli DNA during its replication cycle". Biophysical Journal 1: 337–351. PMC 1366352. PMID 13691706. 
  155. ^ Erica Westly (6 October 2008). "No Nobel for You: Top 10 Nobel Snubs". Scientific American. 
  156. ^ Nobel Prize (1962)
  157. ^ [4]
  158. ^ Wilkins, p. 242
  159. ^ Wilkins, p. 240
  160. ^ Wilkins, p. 243
  161. ^ "The Nobel Prize in Chemistry 1982". Nobel Media AB. Retrieved 21 January 2015. 
  162. ^ Arnott, S.; Kibble, T.W.B.; Shallice, T. (2006). "Maurice Hugh Frederick Wilkins. 15 December 1916 -- 5 October 2004: Elected FRS 1959". Biographical Memoirs of Fellows of the Royal Society 52: 455–478. doi:10.1098/rsbm.2006.0031. PMID 18551798. 
  163. ^ "Iota Sigma Pi professional awards recipients". 2000-07-25. Retrieved 2013-07-25. 
  164. ^ "Biography: Rosalind Franklin". The Royal Borough of Kensington and Chelsea. Retrieved 21 November 2014. 
  165. ^ "Plaque: Rosalind Franklin". London Remembers. Retrieved 27 August 2014. 
  166. ^ "Hampstead Residence". King's College London. Retrieved 21 November 2014. 
  167. ^ "Plaque: Franklin, Gosling, Stokes, Wilson, Wilkins". London Remembers. Retrieved 21 November 2014. 
  168. ^ "The Graduate Houses: Rosalind Franklin Building". Newnham College. Retrieved 27 August 2014. 
  169. ^ Rostvik, Camilla. "Rosalind Franklin's Cambridge, Cambridge, UK". BSHS Travel Guide. Retrieved 27 August 2014. 
  170. ^ Dugard, Jane (18 March 2003). "A grave injustice". Mail & Guardian Online. Retrieved 27 August 2014. 
  171. ^ Sir Aaron Klug opens new Laboratory
  172. ^ "NPG pictures". 1946-06-11. Retrieved 2013-07-25. 
  173. ^ "Dielectrophoresis 2014 Venue and accommodation". Dielectrophoresis 2014. Retrieved 21 November 2014. 
  174. ^ Maddox, p. 323
  175. ^ "seventh annual Rosalind E. Franklin Award for Women in Cancer Research at the National Cancer Institute's Intramural Scientific Retreat [which] honors the commitment of women in cancer research and is given in tribute to chemist Rosalind Franklin, who played a critical role in the discovery of the DNA double helix." The JHU Gazette, Johns Hopkins University, 17 March 2008 For the Record: Cheers
  176. ^ [5]The Royal Society Rosalind Franklin Award (2003): The Royal Society web page. Retrieved 21 July 2006.
  177. ^ "Plaque: DNA at Kings". London Remembers. Retrieved 21 November 2014. 
  178. ^ Dedication[dead link] of Rosalind Franklin University
  179. ^ "Secret of life revisited". Cambridge News. 9 November 2005. Retrieved 1 November 2010. 
  180. ^ "THE ROSALIND FRANKLIN SOCIETY, INC.". NYCorporateList. Retrieved 21 November 2014. 
  181. ^ "Mission". Rosalind Franklin Society. Retrieved 21 November 2014. 
  182. ^ "Rosalind Franklin Fellowships at the Faculty of Mathematics and Natural Sciences, Groningen". European Platform of Women Scientists. 6 October 2014. Retrieved 21 November 2014. 
  183. ^ "€ 6.6 million EU grant for Rosalind Franklin Fellowship". University of Groningen. 2 August 2013. 
  184. ^ "2008 Horwitz Prize Awarded To Arthur Horwich & Ulrich Hartl For Cellular Protein Folding". Medical News Today. 15 October 2008. Retrieved 10 April 2012. 
  185. ^ "Lord Robert Winston opens Rosalind Franklin Building at Nottingham Trent Uni". Nottingham Post (Local World). 16 October 2012. Retrieved 21 November 2014. 
  186. ^ Rosalind Franklin, DNA scientist, celebrated by Google doodle, The Guardian News, 2013, 25 July
  187. ^ "Rosalind Franklin's 93rd Birthday". Google. 
  188. ^ Jonny (10 March 2013). "New plaque for Rosalind Franklin". The Cambridge Tour Company. Retrieved 27 August 2014. 
  189. ^ Ashwell, Louise (10 March 2013). "New plaque unveiled to commemorate unsung heroine of DNA". Varsity (Varsity Publications Ltd.). Retrieved 27 August 2014. 
  190. ^ Kennedy, Caitlin (11 July 2014). "CA Scientist Receives First BIO Rosalind Franklin Award". Biotechnology Industry Organization. 
  191. ^ "Life Story (TV)". Filmaffinity. Retrieved 26 January 2015. 
  192. ^ "Life Story". William Nicholson. Retrieved 26 January 2015. 
  193. ^ "Secret of Photo 51". PBS. Retrieved 26 January 2015. 
  194. ^ "NOVA: DNA - Secret of Photo 51 (2003)". The New York Times. Retrieved 26 January 2015. 
  195. ^ "Secret of Photo 51 Transcript". PBS. Retrieved 26 January 2015. 
  196. ^ "Rosalind Franklin: DNA's Dark Lady (2003) (TV)". MoviesPictures.ORG. Retrieved 4 February 2015. 
  197. ^ "Season 1, Episode 1 The Secret of Life". TVGUIDE. Retrieved 26 January 2015. 
  198. ^ "Episode 1: The Secret of Life". PBS. Retrieved 26 January 2015. 
  199. ^ "Rosalind: A Question of Life". Retrieved 26 January 2015. 
  200. ^ Gearing, Deborah (2006). Burn, and Rosalind: A Question of Life. London: Oberon Books. ISBN 9781840026597. 
  201. ^ Ziegler, Anna (2011). Photograph 51. New York: Dramatists Play Service. ISBN 978-0-8222-2508-9. 
  202. ^ Ziegler, Anna (2011). Photograph 51. New York: Dramatists Play Service. ISBN 978-0-8222-2508-9. 
  203. ^ Grode, Eric (5 November 2010). "The Female Scientist, the Biggest Secret". The New York Times. Retrieved 26 January 2015. 
  204. ^ "Review of ''False Assumptions''". 2013-03-27. Retrieved 2013-07-25. 


Further reading[edit]

  • Brown, Andrew (2007). J.D. Bernal: The Sage of Science. New York: Oxford University Press. ISBN 0-19-920565-5. 
  • Chomet, Seweryn [Ed.] (1995). D.N.A. : Genesis of a Aiscovery. England: Newman-Hemisphere. ISBN 978-1-56700-138-9. 
  • Crick, Francis (1988). What Mad Pursuit : A Personal View of Scientific Discovery. New York: Basic Books. ISBN 0-465-09138-5. 
  • Dickerson, Richard E. (2005). Present at the Flood : How Structural Molecular Biology Came about. Sunderland: Sinauer. ISBN 0-87893-168-6. 
  • Finch, John (2008). A Nobel Fellow on Every Floor : A History of the Medical Research Council Laboratory of Molecular Biology. Cambridge: Medical Research Council Laboratory of Molecular Biology. ISBN 978-1-84046-940-0. 
  • Gibbons, Michelle G. "Reassessing Discovery: Rosalind Franklin, Scientific Visualization, and the Structure of DNA." Philosophy of Science 79 (2012): 63-80.
  • Hager, Thomas (1995). Force of Nature : The Life of Linus Pauling. New York: Simon & Schuster. ISBN 0-684-80909-5. 
  • Horace, Freeland Judson (1996) [1977]. The Eighth Day of Creation: Makers of the Revolution in Biology (Expanded ed.). Plainview, N.Y: CSHL Press. ISBN 0-87969-478-5. 
  • Glynn, Jenifer (2012). My Sister Rosalind Franklin. Oxford: Oxford University Press. ISBN 978-0-19-969962-9. 
  • Glynn, Jenifer (1996). "Rosalind Franklin, 1920-1958". In Shils, Edward. Cambridge Women : Twelve Portraits. Cambridge: Cambridge University Press. pp. 267–282. ISBN 0-521-48287-9. 
  • Klug, Aaron (2004). "R.E. Franklin". In Matthew, H.C.G.; Harrison, Brian. Oxford Dictionary of National Biography : From the Earliest Times to the Year 2000. Oxford: Oxford University Press. ISBN 0-19-861411-X. 
  • Klug, Aaron (2004). "The discovery of the DNA Double Helix". In Krude, Torsten. DNA : Changing Science and Society. Cambridge: Cambridge University Press. pp. 5–27. ISBN 0-52182-378-1. 
  • Olby, Robert (1974). "Rosalind Elsie Franklin". In Gillispie, Charles Coulston. Dictionary of Scientific Biography. V.10. New York: Scribner. ISBN 0-684-10121-1. 
  • Olby, Robert (1994). The Path to The Double Helix : The Discovery of DNA (Unabridged, corrected and enlarged Dover edition ed.). New York: Dover Publications. ISBN 0-486-68117-3. 
  • Olby R (January 2003). "Quiet debut for the double helix". Nature 421 (6921): 402–405. Bibcode:2003Natur.421..402O. doi:10.1038/nature01397. PMID 12540907. 
  • Tait, Sylvia A.S.; Tait, James F. (2004). A Quartet of Unlikely Discoveries. London: Athena Press. ISBN 978-1-8440-1343-2. 
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  • Wilkins, Maurice (2005). The Third Man of the Double Helix : The Autobiography of Maurice Wilkins. Oxford: Oxford Univ. Press. ISBN 978-0-19-280667-3. 
  • Polcovar, Jane (2006). Rosalind Franklin and the Structure of Life. Greensboro, N.C.: Morgan Reynolds Publishing Inc. ISBN 978-1-59935-022-6. 

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