Rosalind Franklin

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

Rosalind Elsie Franklin (25 July 1920 – 16 April 1958)^[1] was an English chemist and X-ray crystallographer who made critical contributions to the understanding of the fine molecular structures of DNA (deoxyribonucleic acid), RNA (ribonucleic acid), viruses, coal, and graphite.^[2] Although her works on coal and viruses were appreciated in her lifetime, her DNA work posthumously achieved the most profound impact as DNA plays a central role in biology, as it carries the genetic information that is passed from parents to their offsprings.

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. She excelled in all major subjects and sports. She passed her matriculation at age eighteen, and won the School Leaving Exhibition of £30 a year for three years. Her father asked her to donate the money she earned to refugee students during World War II. 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, from whom 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 only after two years, due to a disagreeable clash with her director John Randall, and more with her colleague Maurice Wilkins. At Birkbeck, Chair of Physics Department J. D. Bernal 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. According to Francis Crick, her data and research were key in determining the structure.^[4] 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.^[5] Franklin's images of X-ray diffraction, confirming the helical structure of DNA, were imprudently shown to Watson by Wilkins without her permission.^[6]

Her work was published third, in the series of three DNA Nature articles, led by the paper of Watson and Crick.^[7] Watson, Crick and 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.^[8]

After finishing her portion of the work on DNA, with her own research team at Birkbeck College, Franklin led pioneering work on the molecular structures of viruses, including tobacco mosaic virus and the polio virus.^[9] Continuing her research, her team member, and later her beneficiary Aaron Klug went on to win the Nobel Prize in Chemistry in 1982.

Early life and education

Franklin was born in 50 Chepstow Villas,^[10] Notting Hill, London into an affluent and influential British Jewish family.^[11][12] 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 elder brother; Colin (born 1923), Roland (born 1926), and Jenifer (born 1929) were her younger siblings.^[13] 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, known in the family as Mamie, 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] Her uncle, Hugh Franklin, was another prominent figure in the suffrage movement, although his actions therein embarrassed the Franklin family. Her middle name "Elsie" was in memory of Hugh's first wife who died of the First World War flu epidemic.^[13] 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.^[18][19] Franklin's parents helped settle Jewish refugees from Europe who had escaped the Nazis, particularly those from the kindertransport.^[20] They took in two Jewish children to their home, and one of them, a nine-year-old Austrian, Evi Eisenstädter shared Jenifer's room.^[21] (Evi's father Hans Mathias Eisenstädter had been imprisoned in Buchenwald, and after liberation, the family adopted the surname "Ellis".)^[22][23]

From early childhood, Franklin showed exceptional scholastic abilities. At age six, she joined her brother Roland at Norland Place School, a private day school at Holland Park Avenue, 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, & invariably gets her sums right."^[24] 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.^[25] 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,^[26][27] where she excelled in science, Latin,^[28] and sports.^[29] She also learned German, and became fluent in French, the 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 Gustav Holst once called upon her mother to inquire whether she might have suffered from hearing problem or tonsillitis.^[30] With six distinctions, she passed her matriculation in 1938, winning a scholarship for college, the School Leaving Exhibition of £30 a year for three years, and ₤5 from her grandfather.^[31] Her father asked her to give the scholarship to deserving refugee student.^[25]

Cambridge and World War II

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.^[32] In 1941, she was awarded Second Class Honours from her final exams. The distinction was accepted as a bachelor's degree in qualifications for employment. Cambridge began awarding titular B.A. and M.A. degrees to women from 1947, and the previous women graduates retroactively received these.^[33] In her last year at Cambridge, she met a French refugee Adrienne Weill, a former student of Marie Curie, who made a huge influence on her life and career. It was from her that she learned spoken French.^[34]

Franklin was there awarded a research fellowship with which she joined the physical chemistry laboratory of the University of Cambridge to work under Ronald Norrish (winner of the 1967 Nobel Prize in Chemistry), the event described as "without great success."^[35] As described by his biographer, Norrish was "obstinate and almost perverse in argument, overbearing and sensitive to criticism".^[36] He could not decide for her what to work upon, and at that time was succumbing to heavy drinking. Franklin penned that he made her despise him completely.^[37] Resigning from Norrish's Lab, she fulfilled the requirements of the National Service Act and worked as an Assistant Research Officer at the British Coal Utilisation Research Association (BCURA) in 1942.^[9] The BCURA was located on the Coombe Springs Estate near Kingston upon Thames near the southwestern boundary of London. Norrish acted as advisor to the military at BCURA. John G. Bennett was the Director. Marcello Pirani and Victor Goldschmidt, both refugees from the Nazis, were consultants and lectured at BCURA while Franklin attended.^[2] During her BCURA research, she stayed at Adrienne Weill's boarding house in Cambridge till her cousin Irene Franklin asked to join her in a vacated house of her uncle in Putney. With Irene, she volunteered as Air Raid Wardens and regularly made patrols to see the welfare of people during air raids.^[38]

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, she helped classify coals and accurately predict their performance for fuel purposes and for production of wartime devices (i.e. gas masks).^[39] 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'Etat 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).^[40][41]

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. It became part of the mainstream of work on the physics and chemistry of coal, covered by a current monograph,^[42] the annual^[43] and other publications. Mering also continued the study of carbon in various forms, using X-ray diffraction and other methods.^[44]

King's College, London

Double Helix William Astbury Oswald Avery Lawrence Bragg Francis Crick Erwin Chargaff Jerry Donohue Rosalind Franklin Raymond Gosling Frederick Griffith Phoebus Levene Friedrich Miescher Fred Neufeld Sir John Randall Alex Stokes James Watson Maurice Wilkins Herbert Wilson

In 1950, Franklin was granted a three-year Turner and 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.^[45] 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.^[46][47] 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.^[48]

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.^[49] Randall's lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.^[50]

Franklin, working with Gosling,^[51] 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.^[52] When Wilkins inquired about this improved technique, she replied in terms which offended Wilkins as Franklin had "an air of cool superiority".^[53]

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.^[54] 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.^[55][56]

These forms were termed DNA "B" and "A" respectively. Because of the intense personality conflict developing between Franklin and Wilkins, Randall^[57] divided the work on DNA. Franklin chose the data rich A form while Wilkins selected the "B" form^[58][59] 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".^[55]

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.^[60] 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).^[61] During 1952, they worked at applying the Patterson function to the X-ray pictures of DNA they had produced.^[62] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.^[63][64]

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, one day before Crick and Watson had completed their model on the 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.^[65] 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.^[66]

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.^[67] Klug designed this paper to complement the first article he had written defending Franklin's significant contribution to DNA structure.^[68] 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 without her permission.^[69] Watson, in turn, showed Wilkins a prepublication manuscript by Pauling and Corey.^[70] 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 structure

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.^[71] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,^[58][72] 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.^[60][73]

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.^[74]

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".^[75] However, they knew they must complete their model before they could be certain.^[76]

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".^[77] 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.^[78]

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,^[79] 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".^[80]

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.^[5] 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

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 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 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.^[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 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.^[100][101] Casper was not an enthusiastic writer such that 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 Institute of Health, which approved ₤10,000 for three years, the largest fund ever received at Birkbeck.^[103]

The first major international fair after World War II, named Expo 58, was to be held in 1958 in Brussels.^[104][105] 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.^[106] 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.^[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 little 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 at age 27 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 used 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. She 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 John Alfred Ryle, Regius Professor of Physic at the University of Cambridge, as an independent Member of Parliament candidate in 1940, but to no avail.^[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 was then married to Ernst Wohlgemuth,^[23] and had moved to Notting Hill from Chicago, tried matchmaking with Ralph Milliband 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 she 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

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".^[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]} Public Broadcasting Service (PBS) biography even 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 femininist.^[164]

Franklin's letter to her parents in January 1939 is often taken as 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 teacher of science was 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, 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.^[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 (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.^[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. 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."^[5] 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 preceding communication.^[195]

Nobel Prize

Franklin was never nominated for a Nobel Prize.^[196][197] 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.^[198] The first breakthrough was from Matthew Meselson and Franklin Stahl in 1958, who experimentally showed the DNA replication of a bacterium Escherichia coli.^[199] 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,^[200] and of the stepwise chemical reaction.^[201][202] 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.^[4]

In 1962 the Nobel Prize was subsequently awarded to Crick, Watson, and Wilkins.^{[9][203][204]} It is not clear whether she would have been included, had she lived.^[205] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.^[206] 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.^[207] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.^[208] 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."^[209] 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.^[210]

Posthumous recognition

• 1982, Iota Sigma Pi designated Franklin a National Honorary Member.^[211]

• 1992, English Heritage placed a blue plaque on the entrance wall of SW10, Drayton Gardens, 107, Donovan Court, where Rosalind Franklin lived during her professional career until her death.^[212] The inscription reads "Rosalind Franklin, 1920-1958, pioneer of the study of molecular structures including DNA, lived here, 1951-1958".^[213]
• 1993, King's College London renamed the Orchard Residence at their Hampstead Campus on Kidderpore Avenue as Rosalind Franklin Hall.^[214]
• 1993, King's College London placed a blue plaque on its outside wall bearing the inscription: "R. E. Franklin, R. G. Gosling, A. R. Stokes, M. H. F. Wilkins, H. R. Wilson King's/College London/DNA, X-ray, diffraction studies 1953."^[215]
• 1995, Newnham College opened a graduate residence named Rosalind Franklin Building^[216] and put a bust of her in its garden.^[217][218]
• 1997, Birkbeck, University of London School of Crystallography opened the Rosalind Franklin Laboratory.^[219]
• 1997, the asteroid discovered in 1997 was named 9241 Rosfranklin.
• 1998, National Portrait Gallery in London added Rosalind Franklin's portrait next to those of Francis Crick, James Watson and Maurice Wilkins.^[220]
• 1999, the Institute of Physics at Portland Place, London, renamed its theatre as Franklin Lecture Theatre.^[221]
• 2000, King's College London opened the Franklin–Wilkins Building in honour of Franklin's and Wilkins's work at the college.^[222] King's had earlier, in 1994, also named one of the Halls in Hampstead Campus residences in memory of Rosalind Franklin.
• 2001, the American National Cancer Institute established the Rosalind E. Franklin Award for women in cancer research.^[223]
• 2003, the Royal Society established the Rosalind Franklin Award, for an outstanding contribution to any area of natural science, engineering or technology.^[224]
• 2003, the Royal Society of Chemistry declared King's College London as "National Historic Chemical Landmark" and placed a plague on the wall near the entrance of the building, with an epitaph: "Near this site Rosalind Franklin, Maurice Wilkins, Raymond Gosling, Alexander Stokes and Herbert Wilson performed experiments that led to the discovery of the structure of DNA. This work revolutionised our understanding of the chemistry behind life itself."^[225]

• 2004, Finch University of Health Sciences/The Chicago Medical School, located in North Chicago, Illinois, USA changed its name to the Rosalind Franklin University of Medicine and Science.^[226] It also adopted a new motto "Life in Discovery", and Photo 51 as its logo.^[227]
• 2005, the wording on the DNA sculpture (which was donated by James Watson) outside Clare College, Cambridge's Memorial Court is a) on the base: i) "These strands unravel during cell reproduction. Genes are encoded in the sequence of bases." and ii) "The double helix model was supported by the work of Rosalind Franklin and Maurice Wilkins.", as well as b) on the helices: i) "The structure of DNA was discovered in 1953 by Francis Crick and James Watson while Watson lived here at Clare." and ii) "The molecule of DNA has two helical strands that are linked by base pairs Adenine – Thymine or Guanine – Cytosine."^[228]
• 2006, the Rosalind Franklin Society was established in New York,^[229] which aims to recognise, foster, and advance the important contributions of women in the life sciences and affiliated disciplines.^[230]
• 2007, the University of Groningen, supported by the European Union, launched the Rosalind Franklin Fellowship to encourage women researchers to become full university professors.^[231][232]
• 2008, Columbia University awarded an Honorary Louisa Gross Horwitz Prize to Rosalind Franklin, Ph.D., posthumously, "for her seminal contributions to the discovery of the structure of DNA".^[233]
• St Paul's Girls School has established the Rosalind Franklin Technology Centre.^[25]
• 2012, honoured as the namesake of Rosalind, an online project teaching programming via molecular biology.
• 2012, The Right Honourable Professor The Lord Robert Winston opens the Nottingham Trent University multi-million pound research building the Rosalind Franklin Building.^[234]
• 2013, Google honoured Rosalind Franklin with a doodle showing her gazing at a double helix structure of DNA with an X-ray of Photo 51 beyond it.^[235][236]
• 7 March 2013, a plaque was placed on the wall of The Eagle, a pub near Cambridge University where Crick and Watson announced the discovery of the structure of DNA sixty years to the day.^[237][238]
• 2014, BIO Rosalind Franklin Award was established by Biotechnology Industry Organization in collaboration with the Rosalind Franklin Society, for an outstanding woman in the field of industrial biotechnology and bioprocessing.^[239]
• 2014, the Rosalind Franklin University of Medicine and Science unveiled a bronze statue of Franklin, created by Julie Rotblatt-Amrany, near its front entrance.^[240] The ceremony on 29 May was attended by Franklin's nephew Martin Franklin and niece Rosalind Franklin Jekowsky.^[241]

In popular culture

Franklin's part in the discovery of the nature of DNA was shown in the 1987 TV Movie Life Story, starring Tim Pigott-Smith as Crick, Alan Howard as Wilkins, Jeff Goldblum as Watson, and Juliet Stevenson as Franklin. This movie portrayed Franklin as somewhat stern, but also alleged that Watson and Crick did use a lot of her work to do theirs.^[242][243]

A 56-minute documentary of the life and scientific contributions of Franklin, DNA - Secret of Photo 51, was broadcast in 2003 on PBS NOVA.^[244] Narrated by Barbara Flynn, the program features interviews with Wilkins, Gosling, Klug, Maddox,^[245] including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley.^[246] The UK version produced by BBC is titled Rosalind Franklin: DNA's Dark Lady.^[247]

The first episode of another PBS documentary serial, DNA, was aired on 4 January 2004.^[248] 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).^[249]

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,^[250] and published by Oberon Books in 2006.^[251]

Another play, Photograph 51 by Anna Ziegler, published in 2011,^[252] has been produced at several places in the USA.^[253] 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.^[254]

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.^[255]

Publications

Library resources about Rosalind Franklin
Resources in your library Resources in other libraries
By Rosalind Franklin
Resources in your library Resources in other libraries

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

References

Further reading

External links

• The Rosalind Franklin Society
• CWP at UCLA
• The History of medicine topographical database in ub Himetop
• Work at Birbeck and meeting REF (12)
• Work with REF (13)
• REF and the discovery of the structure of DNA. (17)
• REF's death and joining the MRC's LMB in Cambridge (18)
• My aunt, the DNA pioneer by S. Franklin
• "Rosalind Franklin and the Double Helix", Physics Today, by Lynne O. Elkin
• "Light on a Dark Lady", republished article by A. Piper from Trends in Biochemical Science
• Oxford Dictionary of National Biography for article by Sir Aaron Klug
• "The first American newspaper coverage of the discovery of the DNA structure", Saturday, 13 June 1953, The New York Times
• "Biography of Rosalind Franklin", Lynne O. Elkin, Jewish Women Encyclopedia
• Nova: "Secret of Photo 51"
• The Rosalind Franklin Papers Profiles in Science, National Library of Medicine
• Bancroft Archive of the University of California, Berkeley Lynne Elkin collection of original Franklin research
• The Rosalind Franklin Papers at Churchill Archives Centre in Cambridge. The digitised papers are at Codebreakers: Makers of Modern Genetics: the Rosalind Franklin papers
• Rosalind Franklin publications. List of publications from 1948 to (posthumous reprint) 1995
• Key Participants: Rosalind Franklin – Linus Pauling and the Race for DNA: A Documentary History
• "Rosalind Franklin's work on coal, carbon, and graphite", by Peter J F Harris in PDF format
• Energia Vol.6 No.6 (1995) (University of Kentucky centre for Applied Energy Research). Contains Rosalind Franklin Article in PDF format

DNA structure research at King's College London 1947–1959 Rosalind Franklin Raymond Gosling John Randall Alexander Stokes Maurice Wilkins Herbert Wilson

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