Rosalind Franklin

Rosalind Elsie Franklin
Born Rosalind Elsie Franklin
(1920-07-25)25 July 1920
Notting Hill, London, England, United Kingdom
Died 16 April 1958(1958-04-16) (aged 37)
Chelsea, London, England, UK
Ovarian cancer
Resting place Willesden United Synagogue Cemetery
51°32′41″N 0°14′24″W / 51.5447°N 0.2399°W / 51.5447; -0.2399
Citizenship British
Nationality English
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
Virus structures

Rosalind Elsie Franklin (25 July 1920 – 16 April 1958[1]) was an English chemist and X-ray crystallographer who made contributions to the understanding of the molecular structures of DNA (deoxyribonucleic acid), RNA (ribonucleic acid), viruses, coal, and graphite.[2] Although her works on coal and viruses were appreciated in her lifetime, her contributions to the discovery of DNA were largely recognized posthumously.

Born to a prominent British Jewish family, Franklin was educated at a private day school at Norland Place in West London, Lindores School for Young Ladies in Sussex, and St Paul's Girls' School, London. Then she studied the Natural Sciences Tripos at Newnham College, Cambridge, from where she graduated in 1941. Earning a research fellowship, she joined the University of Cambridge physical chemistry laboratory under Ronald George Wreyford Norrish, who disappointed her for his lack of enthusiasm.[3] Fortunately, the British Coal Utilisation Research Association (BCURA) offered her a research position in 1942, and started her work on coals. This helped her earn a PhD in 1945. She went to Paris in 1947 as a chercheur (post-doctoral researcher) under Jacques Mering at the Laboratoire Central des Services Chimiques de l'Etat, where she became an accomplished X-ray crystallographer. She became a research associate at King's College London, in 1951, but was compelled to move to Birkbeck College after two years, owing to disagreeable clashes with her director John Randall and more so with her colleague Maurice Wilkins. At Birkbeck, J. D. Bernal, chair of the physics department, offered her a separate research team. She died in 1958 at the age of 37 of ovarian cancer.

Franklin is best known for her work on the X-ray diffraction images of DNA while at King's College, London, which led to the discovery of the DNA double helix for which James Watson, Francis Crick and Maurice Wilkins shared the Nobel Prize in Physiology or Medicine in 1962. Watson suggested that Franklin would have ideally been awarded a Nobel Prize in Chemistry, along with Wilkins, but the Nobel Committee does not make posthumous nominations.[4][5]

After finishing her work on DNA, Franklin led pioneering work at Birkbeck College, London, on the molecular structures of viruses.[6] Her team member Aaron Klug continued her research, winning the Nobel Prize in Chemistry in 1982.

Early life and education

Franklin was born on 25 July 1920 in 50 Chepstow Villas,[7] Notting Hill, London into an affluent and influential British Jewish family.[8][9] 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. David (born 1919) was the eldest brother; Colin (born 1923), Roland (born 1926), and Jenifer (born 1929) were her younger siblings.[10] 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.[11] Her aunt, Helen Caroline Franklin, known in the family as Mamie, was married to Norman de Mattos Bentwich, who was the Attorney General in the British Mandate of Palestine.[12] Helen Caroline Franklin was active in trade union organisation and the women's suffrage movement, and was later a member of the London County Council.[13][14] Her uncle, Hugh Franklin, was another prominent figure in the suffrage movement, although his actions therein embarrassed the Franklin family. Rosalind's middle name, "Elsie", was in memory of Hugh's first wife, who died in the 1918 flu pandemic.[10] Her family was actively involved with 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.[15][16]

Franklin's parents helped settle Jewish refugees from Europe who had escaped the Nazis, particularly those from the Kindertransport.[17] They took in two Jewish children to their home, and one of them, a nine-year-old Austrian, Evi Eisenstädter, shared Jenifer's room.[18] (Evi's father Hans Mathias Eisenstädter had been imprisoned in Buchenwald, and after liberation, the family adopted the surname "Ellis".)[19][20]

From early childhood, Franklin showed exceptional scholastic abilities. At age six, she joined her brother Roland at Norland Place School, a private day school in West London. At that time, her aunt Mamie (Helen Bentwich), described her to her husband: "Rosalind is alarmingly clever – she spends all her time doing arithmetic for pleasure, and invariably gets her sums right."[21] She also developed an early interest in cricket and hockey. At age nine, she entered a boarding school, Lindores School for Young Ladies in Sussex.[22] The school was near the seaside, and the family wanted a good environment for her delicate health. She was eleven when she went to St Paul's Girls' School, West London, one of the few girls' schools in London that taught physics and chemistry.[22][23][24] At St Paul's she excelled in science, Latin,[25] and sports.[26] She also learned German, and became fluent in French, a language she would later find useful. She topped her classes, and won annual awards. Her only educational weakness was in music, for which the school music director, the composer Gustav Holst, once called upon her mother to inquire whether she might have suffered from hearing problem or tonsillitis.[27] With six distinctions, she passed her matriculation in 1938, winning a scholarship for university, the School Leaving Exhibition of £30 a year for three years, and £5 from her grandfather.[28] Her father asked her to give the scholarship to a deserving refugee student.[22]

Cambridge and World War II

Franklin went to Newnham College, Cambridge in 1938 and studied chemistry within the Natural Sciences Tripos. There she met the spectroscopist Bill Price, who worked with her as a laboratory demonstrator and who later became one of her senior colleagues at King's College London.[29] 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.[30] In her last year at Cambridge, she met a French refugee Adrienne Weill, a former student of Marie Curie, who made a huge influence on her life and career and helped her to improve her spoken French.[31]

Franklin was awarded a research fellowship at Newnham College, with which she joined the physical chemistry laboratory of the University of Cambridge to work under Ronald Norrish, who later won the Nobel Prize in Chemistry. In her one year of work there, she did not have much success.[32] As described by his biographer, Norrish was "obstinate and almost perverse in argument, overbearing and sensitive to criticism".[33] He could not decide for her what to work upon, and at that time was succumbing to heavy drinking. Franklin wrote that he made her despise him completely.[34] Resigning from Norrish's Lab, she fulfilled the requirements of the National Service Acts by working as an assistant research officer at the British Coal Utilisation Research Association (BCURA) in 1942.[6] The BCURA was located on the Coombe Springs Estate near Kingston upon Thames near the southwestern boundary of London. Norrish acted as advisor to the military at BCURA. John G. Bennett was the director. Marcello Pirani and Victor Goldschmidt, both refugees from the Nazis, were consultants and lectured at BCURA while Franklin worked there.[2] During her BCURA research, she stayed at Adrienne Weill's boarding house in Cambridge until her cousin Irene Franklin asked to join her in a vacated house of her uncle in Putney. With Irene, she volunteered as an Air Raid Warden and regularly made patrols to see the welfare of people during air raids.[35]

She studied the porosity of coal using helium to determine its density.[36] Through this, she discovered the relationship between the fine constrictions in the pores of coals and the permeability of the porous space. By concluding that substances were expelled in order of molecular size as temperature increased, she helped classify coals and accurately predict their performance for fuel purposes and for production of wartime devices such as gas masks.[37] 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]

Paris

With World War II ending in 1945, Franklin asked Adrienne Weill for help and to let her know of job openings for "a physical chemist who knows very little physical chemistry, but quite a lot about the holes in coal". At a conference in the autumn of 1946, Weill introduced her to Marcel Mathieu, a director of the Centre national de la recherche scientifique (CNRS), the network of institutes that comprise the major part of the scientific research laboratories supported by the French government. This led to her appointment with Jacques Mering at the Laboratoire Central des Services Chimiques de l'État in Paris. She joined the labo (as referred to by the staff) of Mering on 14 February 1947 as one of the fifteen chercheurs (researchers).[38][39]

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] She published several further papers on this work which has become part of the mainstream of the physics and chemistry of coal. This work was covered in a 1993 monograph,[40] and in the regularly-published textbook Chemistry and Physics of Carbon.[41] Mering continued the study of carbon in various forms, using X-ray diffraction and other methods.[42]

King's College, London

In 1950, Franklin was granted a three-year Turner & Newall Fellowship to work at King's College, London. In January 1951, she started working as a research associate in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall.[43] She was originally appointed to work on X-ray diffraction of proteins and lipids in solution, but Randall redirected her work to DNA fibres because of new developments in the field, and she was to be the only experienced experimental diffraction researcher at King's at the time.[44][45] Randall 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 her.[46]

Even using crude equipment, Wilkins and Gosling had obtained an outstanding diffraction picture of DNA which sparked further interest in this molecule. They had been carrying out X-ray diffraction analysis of DNA in the unit since May 1950, but Randall had not informed them that he had asked Franklin to take over both the DNA diffraction work and guidance of Gosling's thesis.[47] Randall's lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.[48]

Franklin, working with Gosling,[49] started to apply her expertise in X-ray diffraction techniques to the structure of DNA. She used a new fine-focus X-ray tube and microcamera ordered by Wilkins, but which she refined, adjusted and focused carefully. Drawing upon her physical chemistry background, she also skillfully manipulated the critical hydration of her specimens.[50] When Wilkins inquired about this improved technique, she replied in terms which offended Wilkins as Franklin had "an air of cool superiority".[51]

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.[52] 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.[53][54]

Franklin named these two forms "B" and "A" respectively. (The biological functions of A-DNA were only discovered 60 years later.[55]) Because of the intense personality conflict developing between Franklin and Wilkins, Randall[56] divided the work on DNA. Franklin chose the data rich "A" form while Wilkins selected the "B" form[57][58] 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".[53]

By the end of 1951 it was generally accepted at King's that the B form of DNA was a helix, but after she had recorded an asymmetrical image in May 1952, Franklin became unconvinced that the A form of DNA was a helix.[59] In July 1952, as a practical joke on Wilkins (who frequently expressed his view that both forms of DNA were helical), Franklin and Gosling produced a funeral notice regretting the 'death' of helical crystalline DNA (A-DNA).[60] During 1952, they worked at applying the Patterson function to the X-ray pictures of DNA they had produced.[61] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.[62][63]

By January 1953, Franklin had reconciled her conflicting data, concluding that both DNA forms had two helices, and had started to write a series of three draft manuscripts, two of which included a double helical DNA backbone (see below). Her two A-DNA manuscripts reached Acta Crystallographica in Copenhagen on 6 March 1953, one day before Crick and Watson had completed their model on B-DNA. She must have mailed them while the Cambridge team was building their model, and certainly had written them before she knew of their work.[64] 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.[65]

The third draft paper was on the B form of DNA, dated 17 March 1953, which 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.[66] Klug designed this paper to complement the first article he had written defending Franklin's significant contribution to DNA structure.[67] 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.[68] Watson, in turn, showed Wilkins a prepublication manuscript by Pauling and Corey, which contained a DNA structure remarkably like their first incorrect model.[69]

DNA structure

In February 1953, Francis Crick and James 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.[70] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,[57][71] but Franklin was opposed to prematurely building theoretical models, until sufficient data were obtained to properly guide the model building. She took the view that building a model was to be undertaken only after enough of the structure was known.[59][72]

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.[73]

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".[74] However, they knew they must complete their model before they could be certain.[75]

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".[76] This was also one day after Franklin's two A-DNA papers had reached Acta Crystallographica. Wilkins came to see the model the following week, according to Franklin's biographer Brenda Maddox on 12 March, and allegedly informed Gosling on his return to King's.[77]

It is uncertain how long it took for Gosling to inform Franklin at Birkbeck, but her original 17 March B-DNA manuscript does not reflect any knowledge of the Cambridge model. Franklin did modify this draft later before publishing it as the third in the trio of 25 April 1953 Nature articles. On 18 March,[78] 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".[79]

Crick and Watson then published their model in Nature on 25 April 1953 in an article describing the double-helical structure of DNA with only a footnote acknowledging "having been stimulated by a general knowledge of" Franklin and Wilkins' "unpublished" contribution.[80] 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.[81][82]

Weeks later, on 10 April, Franklin wrote to Crick for permission to see their model.[83] 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.[84] 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.[85][86][87]

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 and that she described (in a letter to Adrienne Weill in Paris) as "moving from a palace to the slums... but pleasanter all the same."[88] She was recruited by physics department chair J. D. Bernal,[89] 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).[90] Despite the parting words of Bernal to stop her interest in nucleic acids, she helped Gosling to finish his thesis, although she was no longer his official supervisor. Together they published the first evidence of double helix in the A form of DNA in the 25 July issue of Nature.[91] Moreover, she continued to explore another major nucleic acid, RNA, a molecule equally central to life as DNA. She again used X-ray crystallography to study the structure of the tobacco mosaic virus (TMV), an RNA virus. Her meeting with Aaron Klug in the early 1954 led to 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 all TMV virus particles were of the same length.[92] This was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct.[93]

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.[94] Her colleague Klug worked on spherical viruses with his student John Finch, with Franklin coordinating and overseeing the work.[95] As a team, from 1956 they started publishing seminal works on TMV,[96] cucumber virus 4 and turnip yellow mosaic virus.[97]

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.[98] The Birkbeck team members continued working on RNA viruses affecting several plants, including potato, turnip, tomato and pea.[99] In 1955 the team was joined by an American post-doctoral student Donald Caspar. He worked on the precise location of RNA molecules in TMV. In 1956 he and Franklin published individual but complementary papers in the 10 March issue of Nature, in which they showed that the RNA in TMV is wound along the inner surface of the hollow virus.[100][101] Caspar was not an enthusiastic writer, and Franklin had to write the entire manuscript for him.[102]

In 1957 her research grant from ARC had expired, and was given a one-year extension ending in March 1958. She applied for a new grant from the US National Institutes of Health, which approved £10,000 for three years, the largest fund ever received at Birkbeck.[103]

Expo 58, the first major international fair after World War II, was to be held in Brussels in 1958.[104][105] Franklin was invited to make a five-foot high model of TMV, which she started in 1957. Her materials included table tennis balls and plastic bicycle handlebar grips.[106] The Brussels world's fair, with an exhibit of her virus model at the International Science Pavilion, opened on 17 April, one day after she died.[107]

Her main research team at Birkbeck College, London Klug, Finch and Holmes moved to the Laboratory of Molecular Biology, Cambridge in 1962.[108]

Personal life

Franklin was best described as an agnostic.[109] Her lack of religious faith apparently did not stem from anyone's influence, rather from her own inquisitive mind. She developed her scepticism as a young child. Her mother recalled that she refused to believe in the existence of god, and remarked, "Well, anyhow, how do you know He isn't She?"[110] She later made her position clear, now based on her scientific experience, and wrote to her father in 1940:

[S]cience and everyday life cannot and should not be separated. Science, for me, gives a partial explanation of life... I do not accept your definition of faith i.e. belief in life after death... Your faith rests on the future of yourself and others as individuals, mine in the future and fate of our successors. It seems to me that yours is the more selfish...[111] [as to] 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.[112]

She however did not abandon Jewish traditions. As the only Jewish student at Lindores School, she had Hebrew lessons on her own while her friends went to church.[113] She joined the Jewish Society while in her first term at Newnham College, Cambridge, out of respect of her grandfather's request.[114] She confided to her sister that she was "always consciously a Jew".[112]

Franklin loved travelling abroad, particularly trekking. She first "qualified" at Christmas 1929 for a vacation at Menton, France, where her grandfather went to escape English winter.[115] Her family frequently spent vacations in Wales or Cornwall. A trip to France in 1938 gave her a lasting love for France and its language. She considered the French lifestyle as "vastly superior to that of English".[116] In contrast, she described English people as having "vacant stupid faces and childlike complacency".[117] Her family was almost stuck in Norway in 1939, as World War II was declared on their way home.[118] In another instance, she trekked the French Alps with Jean Kerslake in 1946, which almost cost her her life. She slipped off on a slope, and was barely rescued.[119] But she wrote to her mother, "I am quite sure I could wander happily in France forever. I love the people, the country and the food."[120]

She made several professional trips to US, and was particularly jovial among her American friends and constantly displayed her sense of humour. William Ginoza of the University of California, Los Angeles later recalled that she was the opposite of Watson's description of her, and as Maddox comments, Americans enjoyed her "sunny side."[121]

Watson's The Double Helix almost entirely refers to her as "Rosy", the name people at King's College used behind her back.[122] She did not want to be called by that name because she had a great-aunt Rosy. In the family she was called "Ros".[123] To others, she was simply Rosalind. She made it clear to an American visiting friend Dorothea Raacke, while sitting with her at Crick's table in The Eagle pub in Cambridge. She [Raacke] asked her how she was to be called, and she replied, "I'm afraid it will have to be Rosalind," and added, "Most definitely not Rosy."[124]

She often expressed her political views. She initially blamed Winston Churchill for inciting the war, but later admired him for his speeches. She actively supported Professor John Ryle as an independent candidate for parliament in 1940, but he was unsuccessful.[125]

She did not seem to have intimate relationship with anyone and always kept her deepest personal feelings to herself. Since her younger days she avoided close friendship with the opposite sex. Once her cousins visited them, she paid Roland to accompany them.[116] In her later years, Evi Ellis, who had shared her bedroom when a child refugee and who was then married to Ernst Wohlgemuth[20] and had moved to Notting Hill from Chicago, tried matchmaking with Ralph Miliband but failed. Franklin once told Evi that her flatmate asked her for a drink, but she did not get the intention.[126] She was quite infatuated by her French mentor Mering, who had a wife and a mistress.[120] Mering also admitted that he was captivated by her "intelligence and beauty".[127] According to Sayre, she did confess her feeling for Mering when she was undergoing surgery, but her family denied this. But Mering wept when he visited her later,[125] and destroyed all her letters.[128]

Her closest personal affair was probably with her once post-doctoral student Donald Caspar. In 1956, she visited him at his home in Colorado after her tour to University of California, Berkeley, and she was known to remark later that Caspar was one "she might have loved, might have married". In her letter to Sayre, she described him as "an ideal match".[129]

Illness and death

In mid-1956, while on a work-related trip to the United States, Franklin first began to suspect a health problem. While in New York she found difficulty in zipping her skirt; her stomach had bulged. Back in London she consulted Mair Livingstone, who asked her, "You're not pregnant?" to which she retorted, "I wish I were." But her diagnosis report stated that it was not pregnancy, and her case was marked "URGENT".[130] An operation on 4 September of the same year revealed two tumours in her abdomen.[131] 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,[124] 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.[132] 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.[133]

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.[134] She returned to work in January 1958, and she was given a promotion to Research Associate in Biophysics on 25 February.[135] She fell ill again on 30 March, and she died on 16 April 1958, in Chelsea, London,[136][137] of bronchopneumonia, secondary carcinomatosis, and ovarian cancer. Exposure to X-ray radiation is sometimes considered to be a possible factor in her illness.[138]

Other members of her family have died of cancer, and the incidence of gynaecological cancer is known to be disproportionately high among Ashkenazi Jews.[139] Her death certificate read: A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.[140] She was interred on 17 April 1958 in the family plot at Willesden United Synagogue Cemetery at Beaconsfield Road in London Borough of Brent.[141] The inscription on her tombstone reads:[142][143]

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

Anne Sayre, Franklin's friend and one of her biographers, stated "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".[144] Andrzej Stasiak states "Sayre's book became widely cited in feminist circles for exposing rampant sexism in science."[145] 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".[146] Maddox states, "Randall ... did have many women on his staff ... they found him ... sympathetic and helpful."[147]

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".[148][149] Elkin states that most of the MRC group typically ate lunch together (including Franklin) in the mixed dining room discussed below.[150] 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."[147] Francis Crick also commented that "her colleagues treated men and women scientists alike."[151]

Sayre also discusses at length Franklin's struggle in pursuing science, particularly her father's concern about women in academic professions.[152] This account had been taken to accuse Ellis Franklin of sexism against his daughter. A good deal of information explicitly claims that he strongly opposed her entering Newnham College.[153][154][155][156] Franklin's Public Broadcasting Service (PBS) biography goes further by stating that he refused to pay her fees, and that an aunt stepped in for her.[157] Her sister Jenifer Glynn explains that these stories are myths, and that her parents fully supported Franklin's entire career.[158]

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.[159] His Cambridge colleague, Peter Pauling, wrote in a letter, "Morris [sic] Wilkins is supposed to be doing this work; Miss Franklin is evidently a fool."[160] Crick acknowledges later, "I'm afraid we always used to adopt – let's say, a patronizing attitude towards her."[161]

Glynn accuses Sayre of making her sister a feminist heroine,[162] and Watson's The Double Helix as the root of what she calls "Rosalind Industry". She conjectures that these alleged sexism stories would "have embarrassed her [Rosalind Franklin] almost as much as Watson’s account would have upset her",[158] and declared that "she was never a feminist."[163] Klug and Crick also concurred that she was definitely not a feminist.[164]

Franklin's letter to her parents in January 1939 is often taken as reflecting her own prejudiced attitude and that she was "not immune to the sexism rampant in these circles." In it she remarked one lecturer as "very good, though female."[165] But as Maddox explains, it was more of circumstantial comment rather than a gender bias. It was more of an admiration because at the time woman teachers of science were a rarity. She in fact laughed at men who were embarrassed by the appointment of the first female professor Dorothy Garrod.[166]

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, her position being that the phosphate units are located in the external part of the molecule. She also specified the amount of water to be found in the molecule in accordance with other parts of it, data that have considerable importance in terms of the stability of the molecule. Franklin was the first to discover and formulate these facts, which in fact constituted the basis for all later attempts to build a model of the molecule. However, Watson, at the time ignorant of the chemistry, failed to comprehend the crucial information, and this led to construction of a wrong model.[167]

The other contribution included an X-ray photograph of B-DNA (called photograph 51),[168] that was briefly shown to Watson by Wilkins in January 1953,[169][170] 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.[171][172]

Sayre's biography of Franklin contains a story[173] alleging that the photograph 51 in question was shown to Watson by Wilkins without Franklin's permission,[145][174][175] and that this constituted a case of bad science ethics.[176] Others dispute this story, asserting that Wilkins had been given photograph 51 by Franklin's Ph.D. student Gosling because she was leaving King's to work at Birkbeck, and there was allegedly nothing untoward in this transfer of data to Wilkins[177][178] because Director Randall had insisted that all DNA work belonged exclusively to King's and had instructed Franklin in a letter to even stop working on it and submit her data.[179] Also, it was implied by Horace Freeland Judson, that Maurice Wilkins had taken the photograph out of Franklin's drawer, but this is also said to be incorrect.[180]

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".[181] 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[182] and to post a general statement in Science excusing himself on the basis of being "inexperienced and casual in administrative matters".[183]

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.[184][185]

The Perutz letter was as said one of three letters, published with letters by Wilkins and Watson, which discussed their various contributions. Watson clarified the importance of the data obtained from the MRC report as he had not recorded these data while attending Franklin's lecture in 1951. The upshot of all this was that when Crick and Watson started to build their model in February 1953 they were working with critical parameters that had been determined by Franklin in 1951, and which she and Gosling had significantly refined in 1952, as well as with published data and other very similar data to those available at King's. It was generally believed that Franklin was never aware that her work had been used during construction of the model,[186] but Gosling asserted in his 2013 interview that, "Yes. Oh, she did know about that."[187]

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.[188] Wilkins turned down this offer, as he had taken no part in building the model.[189] 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.[190] There is no doubt that Franklin's experimental data were used by Crick and Watson to build their model of DNA in 1953. Some, including Maddox, have explained this citation omission by suggesting that it may be a question of circumstance, because it would have been very difficult to cite the unpublished work from the MRC report they had seen.[191]

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".[192]

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.[193] 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.[194]

Sayre's early analysis was often ignored because of perceived feminist overtones in her book. Watson and Crick did not cite the X-ray diffraction work of Wilkins and Franklin in their original paper, though they admit having "been stimulated by a knowledge of the general nature of the unpublished experimental results and ideas of Dr. M. H. F. Wilkins, Dr. R. E. Franklin and their co-workers at King's College, London".[80] In fact, Watson and Crick cited no experimental data at all in support of their model. Franklin and Gosling's publication of the DNA X-ray image, in the same issue of Nature, served as the principal evidence:

Thus our general ideas are not inconsistent with the model proposed by Watson and Crick in the preceding communication.[195]

Nobel Prize

Franklin was never nominated for a Nobel Prize.[196][197] Her work was a crucial part in the discovery of DNA, which Francis Crick, James Watson, and Maurice Wilkins were awarded a Nobel Prize in 1962.[198] 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.[199] The first breakthrough was from Matthew Meselson and Franklin Stahl in 1958, who experimentally showed the DNA replication of a bacterium Escherichia coli.[200] 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,[201] and of the stepwise chemical reaction.[202][203] 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.[204]

In 1962 the Nobel Prize was subsequently awarded to Crick, Watson, and Wilkins.[6][205][206] It is not clear whether she would have been included, had she lived.[207] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.[208] 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.[209] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.[210] Watson has suggested that ideally Wilkins and Franklin would have been awarded the Nobel Prize in Chemistry.[4]

Franklin's colleague and principal beneficiary in her will, Aaron 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."[211] This work was exactly what Franklin had started and which she introduced to Klug, and it is highly plausible that, were she alive, she would have shared the Nobel Prize.[212]

Posthumous recognition

Blue plaque on 107 Drayton Gardens, London SW10
Rosalind Franklin University of Medicine and Science at Illinois

Cultural references

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.[251][252]

A 56-minute documentary of the life and scientific contributions of Franklin, DNA - Secret of Photo 51, was broadcast in 2003 on PBS NOVA.[253] Narrated by Barbara Flynn, the program features interviews with Wilkins, Gosling, Klug, Maddox,[254] including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley.[255] The UK version produced by BBC is titled Rosalind Franklin: DNA's Dark Lady.[256]

The first episode of another PBS documentary serial, DNA, was aired on 4 January 2004.[257] 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).[258]

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,[259] and published by Oberon Books in 2006.[260]

Another play, Photograph 51 by Anna Ziegler, published in 2011,[261] has been produced at several places in the USA,[262] and in late 2015 was put on at the Noel Coward Theatre, London, with Nicole Kidman playing Franklin.[263] 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.[264]

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.[265]

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.

Footnotes

  1. ↑ "The Rosalind Franklin Papers, Biographical Information". profiles.nlm.nih.gov. Retrieved 13 November 2011.
  2. 1 2 3 4 5 "The Rosalind Franklin Papers, The Holes in Coal: Research at BCURA and in Paris, 1942–1951". profiles.nlm.nih.gov. Retrieved 13 November 2011.
  3. ↑ Glynn, p. 60
  4. 1 2 "The Discovery of the Molecular Structure of DNA - The Double Helix". Official Website of the Nobel Prizes. Retrieved 4 February 2014.
  5. ↑ "Nobel Prize Facts". Official Website of the Nobel Prizes. Retrieved 24 January 2016.
  6. 1 2 3 "James Watson, Francis Crick, Maurice Wilkins, and Rosalind Franklin". Chemical Heritage Foundation. Retrieved 1 November 2013.
  7. ↑ "Name of Firm: A. Kyeser & Co." (PDF). The Gazette. 22 February 1922. Retrieved 21 November 2014.
  8. ↑ "Rosalind Franklin". London Remembers. Retrieved 21 November 2014.
  9. ↑ GRO Register of Births: SEP 1920 1a 250 KENSINGTON – Rosalind E. Franklin, mmn = Waley
  10. 1 2 Glynn, p.1
  11. ↑ Maddox p. 7
  12. ↑ Segev p.
  13. ↑ Sayre, A. (1975). Rosalind Franklin and DNA. New York: Norton. p. 31. ISBN 0393074935.
  14. ↑ Maddox p. 40
  15. ↑ Maddox, p. 20
  16. ↑ Sayre, p. 35
  17. ↑ Polcovar, p. 20
  18. ↑ Simkin, John (1997). "Rosalind Franklin". Spartacus Educational. Retrieved 13 February 2015.
  19. ↑ "Hans (John) Mathias Eisenstadter Ellis". Geni. Retrieved 13 February 2015.
  20. 1 2 "Evi Ellis". Ancestry.com. Retrieved 13 February 2015.
  21. ↑ Maddox, p. 15
  22. 1 2 3 4 Berger, Doreen (3 December 2014). "A Biography of The Dark Lady Of Notting Hill". United Synagogue Women. Retrieved 7 February 2015.
  23. ↑ Glynn, p. 25
  24. ↑ Sayre p. 41
  25. ↑ Maddox p. 30
  26. ↑ Maddox, p. 26
  27. ↑ Glynn, p. 28
  28. ↑ Glynn, p. 30
  29. ↑ Dixon, R. N.; Agar, D. M.; Burge, R. E. (1997). "William Charles Price. 1 April 1909−−10 March 1993". Biographical Memoirs of Fellows of the Royal Society 43: 438. doi:10.1098/rsbm.1997.0023. JSTOR 770344.
  30. ↑ Fact sheet: Women at Cambridge: A Chronology, . Archived 14 January 2012 at the Wayback Machine.
  31. ↑ Polcovar, p. 31
  32. ↑ Rosalind Franklin, Cold Spring Harbor Laboratory's Dolan DNA Learning Center, ID 1649, .
  33. ↑ Dainton, F.; Thrush, B. A. (1981). "Ronald George Wreyford Norrish. 9 November 1897-7 June 1978". Biographical Memoirs of Fellows of the Royal Society 27: 379–424. doi:10.1098/rsbm.1981.0016. JSTOR 769878.
  34. ↑ Maddox, p. 72
  35. ↑ Polcovar, p. 37
  36. ↑ Harris, P.J.F. (March 2001). "Rosalind Franklin's work on coal, carbon, and graphite" (PDF). Interdisciplinary Science Reviews 26 (3): 204–210. doi:10.1179/030801801679467.
  37. ↑ "The Rosalind Franklin Papers: The Holes in Coal: Research at BCURA and in Paris, 1942–1951". Profiles.nlm.nih.gov. Retrieved 2013-07-25.
  38. ↑ "Rosalind Franklin". Timetoast. Retrieved 28 August 2014.
  39. ↑ "Rosalind Franklin (1920–1958)". DNA Learning Center, Cold Spring Harbor Laboratory. Retrieved 28 August 2014.
  40. ↑ D. W. van Krevelen, Coal, Third Edition: Typology – Physics – Chemistry – Constitution, Elsevier, New York, 1993.
  41. ↑ Chemistry and Physics of Carbon, vol 1–, 1968–, Elsevier, New York.
  42. ↑ G. Terriere, A. Oberlin, J. Mering, Oxidation of graphite in liquid medium -- observations by means of microscopy and electron diffraction, Carbon, 5, 431--, 1967.
  43. ↑ Maddox, p. 124
  44. ↑ Maddox, p. 114
  45. ↑ Wilkins, Wilkins, M., The Third Man of the Double Helix, an autobiography (2003) Oxford University Press, Oxford. pp. 143–144
  46. ↑ Wilkins, p. 121
  47. ↑ 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).
  48. ↑ Sayre, Olby, Maddox, Elkin, Wilkins
  49. ↑ Maddox, p. 129
  50. ↑ Elkin, p. 43
  51. ↑ Wilkins p. 155
  52. ↑ Elkin p. 45
  53. 1 2 Maddox, p. 153
  54. ↑ Wilkins, p. 154
  55. ↑ Wood, Bayden R. (2016). "The importance of hydration and DNA conformation in interpreting infrared spectra of cells and tissues". Chemical Society Reviews. Online. doi:10.1039/C5CS00511F. PMID 26403652.
  56. ↑ Maddox p 155
  57. 1 2 Wilkins, p. 158
  58. ↑ Maddox, p. 155
  59. 1 2 Wilkins, p. 176
  60. ↑ Wilkins, p. 182
  61. ↑ Maddox, p. 168
  62. ↑ Maddox, p. 169
  63. ↑ Wilkins, pp. 232–233
  64. ↑ Maddox p. 199
  65. ↑ Franklin and Gosling (1953). Acta Crystallographica, 6, 673-677
  66. ↑ Klug, A. "Rosalind Franklin and the Double Helix", Nature 248 (26 April 1974): 787–788
  67. ↑ Klug, A. Rosalind Franklin and the Discovery of the Structure of DNA, Nature 219 (24 August 1968): 808–810 & 843.
  68. ↑ Hubbard, Ruth (2013). "Science, Power, Gender: How DNA Became the Book of Life". Women, Science, and Technology (3rd ed.). Hoboken: Taylor and Francis. p. 269. ISBN 1-135-05542-4.
  69. ↑ Yockey, pp. 9–10
  70. ↑ Crick's 31 December 1961 letter to Jacque Monod cited above
  71. ↑ Maddox, p. 147
  72. ↑ Maddox, p. 161
  73. ↑ Hubbard, Ruth (1990). The Politics of Women's Biology. Rutgers State University. p. 60. ISBN 0-8135-1490-8.
  74. ↑ "The Double Helix" p. 115
  75. ↑ "The Double Helix" p. 60
  76. ↑ "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.
  77. ↑ Maddox p. 207
  78. ↑ In contrast to his other letters to Crick, Wilkins dated this one.
  79. ↑ "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.
  80. 1 2 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. Watson and Crick's article was immediately followed by the two King's College submissions: Wilkins MHF, Stokes AR, Wilson HR (April 1953). "Molecular structure of deoxypentose nucleic acids" (PDF). 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.
  81. ↑ Franklin and Gosling (1953)
  82. ↑ Maddox, p. 210
  83. ↑ 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.
  84. ↑ Holt, J. (2002)
  85. ↑ 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.
  86. ↑ Scher, Stanley (2004). "Was Watson and Crick's model truly self-evident?". Nature 427 (6975): 584–584. Bibcode:2004Natur.427..584S. doi:10.1038/427584c. PMID 14961092.
  87. ↑ 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.
  88. ↑ Maddox, p. 205
  89. ↑ Maddox, p. 229
  90. ↑ Maddox, p. 235
  91. ↑ Franklin, RE; Gosling, RG (1953). "Evidence for 2-chain helix in crystalline structure of sodium deoxyribonucleate.". Nature 172 (4369): 156–157. Bibcode:1953Natur.172..156F. doi:10.1038/172156a0. PMID 13072614.
  92. ↑ Franklin, RE. (1955). "Structure of Tobacco Mosaic Virus". Nature 175 (4452): 379–381. Bibcode:1955Natur.175..379F. doi:10.1038/175379a0. PMID 14356181.
  93. ↑ Maddox, p. 252
  94. ↑ Franklin and Holmes, 1956
  95. ↑ Maddox, p. 254
  96. ↑ 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 (3): 403–416. doi:10.1016/0006-3002(56)90463-2. PMID 13315300.
  97. ↑ Franklin et al., 1958
  98. ↑ Maddox, p. 256
  99. ↑ Maddox, p. 262
  100. ↑ Franklin, RE (1956). "Structure of Tobacco Mosaic Virus: Location of the Ribonucleic Acid in the Tobacco Mosaic Virus Particle". Nature 177 (4516): 928–930. Bibcode:1956Natur.177..928F. doi:10.1038/177928b0.
  101. ↑ Casper, D. L. D. (1956). "Structure of Tobacco Mosaic Virus: Radial Density Distribution in the Tobacco Mosaic Virus Particle". Nature 177 (4516): 928–928. Bibcode:1956Natur.177..928C. doi:10.1038/177928a0.
  102. ↑ Maddox, p. 269
  103. ↑ Glynn, p. 145
  104. ↑ "Expo 58". Retrieved 21 January 2015.
  105. ↑ Devos, Rika (2011). "Expo 58: the catalyst for Belgium's Welfare State Government complex?". Planning Perspectives 26 (4): 649–659. doi:10.1080/02665433.2011.599934.
  106. ↑ "Behind the picture: Rosalind Franklin and the polio model". Medical Research Council. Retrieved 21 January 2015.
  107. ↑ Maddox, Brenda. "Mother of DNA". New Humanist. Retrieved 21 January 2015.
  108. ↑ Glynn, p. 153
  109. ↑ "Rosalind Franklin". NNDB. Soylent Communications. Retrieved 8 February 2015.
  110. ↑ Glynn, p. 12
  111. ↑ Glynn, p. 62
  112. 1 2 Maddox, p. 61
  113. ↑ Glynn, p. 19
  114. ↑ Glynn, p. 44
  115. ↑ Glynn, p. 16
  116. 1 2 Polcovar, p. 33
  117. ↑ Polcovar, p. 59
  118. ↑ Glynn, p. 33
  119. ↑ Glynn, p. 79
  120. 1 2 Polcovar, p. 41
  121. ↑ Maddox, p. 277
  122. ↑ Watson, p. 16
  123. ↑ Glynn, p. 157
  124. 1 2 Maddox, p. 288
  125. 1 2 Glynn, p. 52
  126. ↑ Maddox, p. 261
  127. ↑ Polcovar, p. 51
  128. ↑ Maddox, p. 287
  129. ↑ Maddox, p. 283
  130. ↑ Maddox, p. 284
  131. ↑ Maddox, p. 285
  132. ↑ Maddox, p. 292
  133. ↑ Maddox, p. 296
  134. ↑ Maddox, p. 301
  135. ↑ Maddox, p. 302
  136. ↑ GRO Register of Deaths: JUN 1958 5c 257 CHELSEA – Rosalind E. Franklin, age 37
  137. ↑ Maddox, pp. 305–307
  138. ↑ "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
  139. ↑ Maddox, p.320
  140. ↑ Murray, Ruby J (2011). "Historical Profile: Rosalind Franklin". Dumbo Feather. Retrieved 27 August 2014.
  141. ↑ "Rosalind Elsie Franklin". Find A Grave. 19 October 2001. Retrieved 27 August 2014.
  142. ↑ "Rosalind Franklin tomb". Himetop. Retrieved 27 August 2014.
  143. ↑ Friedman, Meyer; Friedland, Gerald W. Medicine'S 10 Greatest Discoveries. Universities Press. p. 227. ISBN 9788173712265.
  144. ↑ Sayre, p 96
  145. 1 2 Stasiak, Andrzej (March 2001). "Rosalind Franklin". EMBO Reports 2 (3): 181. doi:10.1093/embo-reports/kve037. PMC 1083834.
  146. ↑ Hussain, Farooq (20 November 1975). Did Rosalind Franklin deserve DNA Nobel prize?. New Scientist 68 (Reed Business Information). p. 470. Retrieved 10 January 2011.
  147. 1 2 Maddox, p. 135
  148. ↑ Sayre, p.97
  149. ↑ Bryson, B. (2004) p. 490
  150. ↑ Elkin, p.45
  151. ↑ Crick, p. 68
  152. ↑ Sayre, pp. 42–45
  153. ↑ McGrayne, p. 6
  154. ↑ "Rosalind Franklin". The Manhattan Project Heritage Preservation Association, Inc. Retrieved 13 February 2015.
  155. ↑ Lewis, Jone Johnson. "Rosalind Franklin". About Education. Retrieved 13 February 2015.
  156. ↑ "Rosalind Franklin". What is Biotechnology. Retrieved 13 February 2015.
  157. ↑ "Rosalind Franklin". PBS Online. Retrieved 13 February 2015.
  158. 1 2 Glynn, Jenifer (2012). "Remembering my sister Rosalind Franklin". The Lancet 379 (9821): 1094–1095. doi:10.1016/S0140-6736(12)60452-8. PMID 22451966.
  159. ↑ 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.
  160. ↑ "Quotes by or related to Rosalind Franklin". Oregon State University Libraries. Retrieved 14 February 2015.
  161. ↑ McGrayne, p. 318
  162. ↑ Glynn, J. (2008). "Rosalind Franklin: 50 years on". Notes and Records of the Royal Society 62 (2): 253–255. doi:10.1098/rsnr.2007.0052.
  163. ↑ Glynn, p. 158
  164. ↑ Crick, p. 69
  165. ↑ 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
  166. ↑ Maddox, p. 48
  167. ↑ Everson, Ted (2007). The Gene: A Historical Perspective. Westport, Connecticut: Greenwood Press. p. 85. ISBN 978-0-313-33449-8.
  168. ↑ Maddox, pp. 177–178
  169. ↑ Maddox, p. 196
  170. ↑ Crick, (1988) p. 67.
  171. ↑ Elkin, L.O. (2003)p 44
  172. ↑ Maddox, pp. 198–199
  173. ↑ Sayre, p. 151
  174. ↑ Minkoff, Eli; Baker, Pamela (2000). Biology Today: An Issues Approach (2 ed.). New York: Garland Publishing. p. 58. ISBN 978-0-8153-2760-8.
  175. ↑ Creager, Angela (2003). "Crystallizing a Life in Science". American Scientist. Sigma Xi, The Scientific Research Society. Retrieved 25 January 2015.
  176. ↑ Stasiak, Andrzej (2003). "The First Lady of DNA". EMBO Reports 4 (1): 14. doi:10.1038/sj.embor.embor723. PMC 1315822.
  177. ↑ Maddox, pp. 196
  178. ↑ Wilkins, p. 198
  179. ↑ Maddox p.312,
  180. ↑ Wilkins, p. 257
  181. ↑ Maddox p.188
  182. ↑ 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.
  183. ↑ Science, 27 June 1969, pp. 207–212, also reprinted in the Norton critical edition of The Double Helix, edited by Gunther Stent.
  184. ↑ Maddox, p. 199
  185. ↑ Watson (1969).
  186. ↑ Maddox, p. 316
  187. ↑ 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.
  188. ↑ Wilkins, p. 213
  189. ↑ Wilkins, p. 214
  190. ↑ Wilkins, p. 226
  191. ↑ Maddox, p. 207
  192. ↑ Maddox, pp316–317, and other parts of the epilogue
  193. ↑ Watson, J.D. (1968) pp. 95–96
  194. ↑ Sayre,A. (1975) pp. 156–167
  195. ↑ 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.
  196. ↑ "The Discovery of the Molecular Structure of DNA – The Double Helix". Nobelprize.org. 2003-09-30. Retrieved 2013-07-25.
  197. ↑ Washington, Harriet A. (2012-12-31). "Don't Forget Rosalind Franklin". Ms.
  198. ↑ Beard, Mary (2001). "Down among the Women (Nobel Laureates)". The Kenyon Review (Harvard University Press) 23 (2): 239–247. JSTOR 4338226.
  199. ↑ Gann, Alexander; Witkowski, Jan A. (2013). "DNA: Archives reveal Nobel nominations". Nature 496 (7446): 434. Bibcode:2013Natur.496..434G. doi:10.1038/496434a. PMID 23619686.
  200. ↑ Meselson, Matthew; Stahl, Franklin W. (1958). "The replication of DNA in Escherichia coli". Proceedings of the National Academy of Sciences 44 (7): 671–682. Bibcode:1958PNAS...44..671M. doi:10.1073/pnas.44.7.671. PMC 528642. PMID 16590258.
  201. ↑ Nakada, D; Ryan, FJ (1961). "Replication of deoxyribonucleic acid in non-dividing bacteria". Nature 189 (4762): 398–399. Bibcode:1961Natur.189..398N. doi:10.1038/189398a0. PMID 13727575.
  202. ↑ 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. doi:10.1002/jcp.1030570107. PMID 13724093.
  203. ↑ 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. Bibcode:1961BpJ.....1..337C. doi:10.1016/S0006-3495(61)86893-8. PMC 1366352. PMID 13691706.
  204. ↑ 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
  205. ↑ Erica Westly (6 October 2008). "No Nobel for You: Top 10 Nobel Snubs". Scientific American.
  206. ↑ Nobel Prize (1962)
  207. ↑ Hartocollis, Anemona (3 December 2014). "By Selling Prize, a DNA Pioneer Seeks Redemption". The New York Times. Retrieved 13 February 2015.
  208. ↑ "The Nobel Prize in Physiology or Medicine 1962". Nobel Media AB. Retrieved 13 February 2015.
  209. ↑ Wilkins, p. 240
  210. ↑ Wilkins, p. 243
  211. ↑ "The Nobel Prize in Chemistry 1982". Nobelprize.org. Nobel Media AB. Retrieved 21 January 2015.
  212. ↑ 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.
  213. ↑ "Iota Sigma Pi professional awards recipients". Iotasigmapi.info. 2000-07-25. Retrieved 2013-07-25.
  214. ↑ "Biography: Rosalind Franklin". The Royal Borough of Kensington and Chelsea. Retrieved 21 November 2014.
  215. ↑ "Plaque: Rosalind Franklin". London Remembers. Retrieved 27 August 2014.
  216. ↑ "Hampstead Residence". King's College London. Retrieved 21 November 2014.
  217. ↑ "Plaque: Franklin, Gosling, Stokes, Wilson, Wilkins". London Remembers. Retrieved 21 November 2014.
  218. ↑ "The Graduate Houses: Rosalind Franklin Building". Newnham College. Retrieved 27 August 2014.
  219. ↑ Rostvik, Camilla. "Rosalind Franklin's Cambridge, Cambridge, UK". BSHS Travel Guide. Retrieved 27 August 2014.
  220. ↑ Dugard, Jane (18 March 2003). "A grave injustice". Mail & Guardian Online. Retrieved 27 August 2014.
  221. ↑ Sir Aaron Klug opens new Laboratory
  222. ↑ "NPG pictures". Npg.org.uk. 1946-06-11. Retrieved 2013-07-25.
  223. ↑ "Dielectrophoresis 2014 Venue and accommodation". Dielectrophoresis 2014. Retrieved 21 November 2014.
  224. ↑ "The future". King's College London. Retrieved 12 May 2015.
  225. ↑ "Rosalind E. Franklin Award". National Cancer Institute. Retrieved 14 February 2015.
  226. ↑ "Rosalind Franklin Fellowships at the Faculty of Mathematics and Natural Sciences, Groningen". European Platform of Women Scientists. 6 October 2014. Retrieved 21 November 2014.
  227. ↑ "€ 6.6 million EU grant for Rosalind Franklin Fellowship". University of Groningen. 2 August 2013.
  228. ↑ The Royal Society Rosalind Franklin Award (2003): The Royal Society web page. Retrieved 21 July 2006.
  229. ↑ "Plaque: DNA at Kings". London Remembers. Retrieved 21 November 2014.
  230. ↑ "History". Rosalind Franklin University of Medicine and Science. Retrieved 13 February 2015.
  231. ↑ "University Honors Namesake With New Sculpture". Rosalind Franklin University of Medicine and Science. Retrieved 13 February 2015.
  232. ↑ "2016 Rosalind Franklin Young Investigator Award of The Gruber Foundation". Genetics Society of America. Retrieved 25 October 2015.
  233. ↑ "Secret of life revisited". Cambridge News. 9 November 2005. Retrieved 1 November 2010.
  234. ↑ "THE ROSALIND FRANKLIN SOCIETY, INC.". NYCorporateList. Retrieved 21 November 2014.
  235. ↑ "Mission". Rosalind Franklin Society. Retrieved 21 November 2014.
  236. ↑ "2008 Horwitz Prize Awarded To Arthur Horwich & Ulrich Hartl For Cellular Protein Folding". Medical News Today. 15 October 2008. Retrieved 10 April 2012.
  237. ↑ "Researchers Launch Innovative, Hands-on Online Tool for Science Education". Jacobsschool.ucsd.edu. Retrieved 14 January 2016.
  238. ↑ "Lord Robert Winston opens Rosalind Franklin Building at Nottingham Trent Uni". Nottingham Post (Local World). 16 October 2012. Retrieved 21 November 2014.
  239. ↑ Rosalind Franklin, DNA scientist, celebrated by Google doodle, The Guardian News, 2013, 25 July
  240. ↑ "Rosalind Franklin's 93rd Birthday". Google.
  241. ↑ Jonny (10 March 2013). "New plaque for Rosalind Franklin". The Cambridge Tour Company. Retrieved 27 August 2014.
  242. ↑ Ashwell, Louise (10 March 2013). "New plaque unveiled to commemorate unsung heroine of DNA". Varsity (Varsity Publications Ltd.). Retrieved 27 August 2014.
  243. ↑ Kennedy, Caitlin (11 July 2014). "CA Scientist Receives First BIO Rosalind Franklin Award". Biotechnology Industry Organization.
  244. ↑ "Rosalind Franklin University unveils bronze statue of its namesake". Daily Herald. 29 May 2014. Retrieved 13 February 2015.
  245. ↑ "Franklin STEM Elementary". Schoolwires, Inc. Retrieved 25 October 2015.
  246. ↑ TCAJOBS (14 October 2014). "Building Tri-Cities: Rosalind Franklin STEM Elementary". Tri-Cities Area Journal of Business. Retrieved 25 October 2015.
  247. ↑ "The Rosalind Franklin Science Building". University of Wolverhampton. Retrieved 25 October 2015.
  248. ↑ "£25m lab block for the University of Wolverhampton". Express & Star. 17 October 2014. Retrieved 25 October 2015.
  249. ↑ "Boats - Newnham College Boat Club". Retrieved 5 November 2015.
  250. ↑ "» Professor Dame Anne Glover to deliver inaugural Rosalind Franklin Lecture on 10 March". British Humanist Association. Retrieved 2016-03-07.
  251. ↑ "Life Story (TV)". Filmaffinity. Retrieved 26 January 2015.
  252. ↑ "Life Story". William Nicholson. Retrieved 26 January 2015.
  253. ↑ "Secret of Photo 51". PBS. Retrieved 26 January 2015.
  254. ↑ "NOVA: DNA - Secret of Photo 51 (2003)". The New York Times. Retrieved 26 January 2015.
  255. ↑ "Secret of Photo 51 Transcript". PBS. Retrieved 26 January 2015.
  256. ↑ "Rosalind Franklin: DNA's Dark Lady (2003) (TV)". MoviesPictures.ORG. Retrieved 4 February 2015.
  257. ↑ "Season 1, Episode 1 The Secret of Life". TVGUIDE. Retrieved 26 January 2015.
  258. ↑ "Episode 1: The Secret of Life". PBS. Retrieved 26 January 2015.
  259. ↑ "Rosalind: A Question of Life". deborahgearing-playwright.moonfruit.com. Retrieved 26 January 2015.
  260. ↑ Gearing, Deborah (2006). Burn, and Rosalind: A Question of Life. London: Oberon Books. ISBN 9781840026597.
  261. ↑ Ziegler, Anna (2011). Photograph 51. New York: Dramatists Play Service. ISBN 978-0-8222-2508-9.
  262. ↑ Ziegler, Anna (2011). Photograph 51. New York: Dramatists Play Service. ISBN 978-0-8222-2508-9.
  263. ↑ Porteous, Jacob (24 April 2015). "Nicole Kidman Returns To The West End In Photograph 51". London Theatre Direct. Retrieved 27 October 2015.
  264. ↑ Grode, Eric (5 November 2010). "The Female Scientist, the Biggest Secret". The New York Times. Retrieved 26 January 2015.
  265. ↑ "Review of ''False Assumptions''". Productionottawa.com. 2013-03-27. Retrieved 2013-07-25.

References

Further reading

  • 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 (2012). "Reassessing Discovery: Rosalind Franklin, Scientific Visualization, and the Structure of DNA". Philosophy of Science 79: 63–80. doi:10.1086/663241. 
  • 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 (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 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. 
  • 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. 

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