Rosalind Franklin

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Rosalind Franklin
Rosalind Franklin
Rosalind Franklin
Born 25 July 1920 (1920-07-25)
Notting Hill, London, United Kingdom
Died 16 April 1958 (1958-04-17) (age 37)
Chelsea, London, United Kingdom
Cancer of the Ovary
Nationality UK citizen
Fields 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
Known for Fine structure of coal and graphite, DNA structure, viruses
Religious stance From Jewish family but personal beliefs unknown

Rosalind Elsie Franklin (25 July 1920 Notting Hill, London – 16 April 1958 Chelsea, London) was an English biophysicist and X-ray crystallographer who made important contributions to the understanding of the fine molecular structures of DNA, viruses, coal and graphite. Franklin is still best known for her work on the X-ray diffraction images of DNA. Her data, according to Francis Crick, were “the data we actually used” [1] to formulate Crick and Watson's 1953 hypothesis regarding the structure of DNA.[2] However, when her work was published third, in the series of three DNA Nature articles, it appeared to only represent evidence in support of their hypothesis.[3] The possibility of Franklin having played a major role was not revealed until Watson wrote his personal account, The Double Helix,[4] which subsequently inspired several people to investigate DNA history and Franklin’s contribution. The first, Robert Olby’s "The Path to the Double Helix," supplied information about original source materials for those that followed.[5] After finishing her portion of the DNA work, Franklin led pioneering work on the tobacco mosaic and polio viruses. She died aged 37 of complications arising from ovarian cancer.


[edit] Background

Franklin was born in Notting Hill, London[6] into an affluent and influential British-Jewish family.[7] Her father was Ellis Arthur Franklin (1894-1964), a London merchant banker and her mother was Muriel Frances Waley (1894-1976); she was the elder daughter and second of the family of five children.

Her uncle was Herbert Samuel (later Viscount Samuel) who was Home Secretary in 1916 and the first practicing Jew to serve in the British Cabinet.[8] He was also the first High Commissioner (effectively governor) for the British Mandate of Palestine.

Her aunt Helen Carolin Franklin was married to Norman de Mattos Bentwich, who was Attorney General in the British Mandate of Palestine.[9] She was active in trade union organisation and women's suffrage, and was later a member of the London County Council.[10][11]

Franklin was educated at St Paul's Girls' School[12][13] where she excelled in science, Latin[14] and sport.[15] Her family were actively involved with a Working Men's College, where Ellis Franklin, her father, taught electricity, magnetism and the history of the Great War in the evenings and later became vice principal.[16][17] Later Franklin's family helped settle Jewish refugees from Europe who had escaped the Nazis.[11]

In the summer of 1938 Franklin went to Newnham College, Cambridge. She passed her finals in 1941, but was only awarded a degree titular, as women were not entitled to degrees (BA Cantab.) from Cambridge at the time; in 1945 Franklin received her PhD from Cambridge University.

[edit] British Coal Utilisation Research Association

Franklin worked for Ronald Norrish between 1941 and 1942. Because of her desire to contribute to the World War II effort, she worked at the British Coal Utilisation Research Association in Kingston-upon-Thames from August 1942, studying the porosity of coal. Her work helped spark the idea of high-strength carbon fibres and was the basis of her 1945 doctoral thesis - "The physical chemistry of solid organic colloids with special reference to coal and related materials." [18][19]

[edit] Laboratoire central des services chimiques de l'État

After the war ended Franklin accepted an offer to work in Paris with Jacques Mering.[20] She learned x-ray diffraction techniques on coal and related inorganic materials during her three years at the Laboratoire central des services chimiques de l'État,[21]but she never fully trained as a crystallographer working on organic chemicals. This later proved to be a problem for her DNA work. Franklin seemed to have been very happy there[22] and earned an international reputation based on her published research on the structure of coal.[23] In 1950 she sought work in England[24] and in June 1950 she was appointed to a position at King's College London.[25]

[edit] King's College London

Double Helix
William Astbury
Oswald Avery
Francis Crick
Erwin Chargaff
Max Delbrück
Jerry Donohue
Rosalind Franklin
Raymond Gosling
Phoebus Levene
Linus Pauling
Sir John Randall
Erwin Schrödinger
Alex Stokes
James Watson
Maurice Wilkins
Herbert Wilson

In January 1951, Franklin started working as a research associate at King's College London in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall.[26] Although originally she was to have worked on x-ray diffraction of proteins and lipids in solution, Randall redirected her work to DNA fibers before she started working at King's since Franklin was to be the only experienced experimental diffraction researcher at King’s in 1951.[27][28] He made this reassignment, even before she started working at King's, because of the following pioneering work by Maurice Wilkins and Raymond Gosling.[29] Even using crude equipment these two men had obtained an outstanding diffraction picture of DNA which sparked further interest in this molecule. Wilkins and Gosling had been carrying out x-ray diffraction analysis of DNA in the Unit since May 1950, but Randall had not informed them of his having asked Franklin to take over both the DNA diffraction work and guidance of Gosling’s thesis.[30] Randall’s lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.[31] Franklin, working with her student Raymond Gosling,[32] started to apply her expertise in x-ray diffraction techniques to the structure of DNA. She used a new fine focus X-ray tube and microcamera ordered by Wilkins, but which she refined, adjusted and focused carefully. Drawing upon her physical chemistry background, Franklin also skillfully manipulated the critical hydration of her specimens.[33] When Wilkins inquired about this improved technique, Franklin replied in terms which offended Wilkins as Franklin having “an air of cool superiority.” [34]Franklin’s habit of intensely looking people in the eye while being concise, impatient and directly confrontational to the point of abrasiveness 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. [35] In spite of the intense atmosphere, Franklin and Gosling discovered that there were two forms of DNA: at high humidity (when wet) the DNA fiber became long and thin, when it was dried it became short and fat.[36][37] These were termed DNA 'B' and 'A' respectively. Because of the intense personality conflict developing between Franklin and Wilkins, Randall [38] divided the work on DNA. Franklin chose the data rich A form while Wilkins selected the 'B' form[39][40] 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, "amongst the most beautiful x-ray photographs of any substance ever taken".[36]

By the end of 1951 it was generally accepted in King's that the B form of DNA was a helix, but Franklin became unconvinced that the A form of DNA was helical in structure[41]after she had recorded an asymmetrical image in 1952 May. As a practical joke on Wilkins (who frequently expressed his view that DNA was helical), Franklin and Gosling produced a death notice regretting the 'death' of helical crystalline DNA (A-DNA).[42] During 1952 Rosalind Franklin and Raymond Gosling worked at applying the Patterson function to the x-ray pictures of DNA they had produced.[43] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.[44][45]

Franklin and Gosling death notice for a helical structure for crystalline DNA (or A-DNA)

By 1953 January, Franklin had reconciled her conflicting data 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,[46] one day before Crick and Watson had completed their model.[47] Franklin had to have mailed them while the Cambridge team was building their model, and certainly had written them before she knew of their work. On 8 July, 1953 she modified one of these “in proof,” Acta articles “in light of recent work” by the King’s and Cambridge research teams.[48] The third draft paper on the ‘B’ form of DNA, dated 17 January 1953, was discovered years later amongst her papers, by Franklin’s Birkbeck colleague, Aaron Klug. He then published an evaluation of the draft’s close correlation with the third of the original trio of 25 April 1953 Nature DNA articles.[49]Klug designed this paper to complement the first article he had written defending Franklin’s significant contribution to DNA structure.[50]. 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 traveled 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. Wilkin commiserated with his harried friend and then changed the course of DNA history with the following disclosure. Watson was shown (by Wilkins) Franklin's famous photograph 51, which had been given to Wilkins by Gosling. Watson, in turn, showed Wilkins a pre-publication manuscript by Pauling and Corey.[51] 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.

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 similar data 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, with Franklin’s being the most unique and critical data completed by February 1953.[52] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,[39][53] but Franklin was opposed to prematurely building theoretical models,until sufficient data was obtained to properly guide the model building. She took the view that building a model was only to be undertaken after enough of the structure was known.[41][54]Ever cautious she wanted to eliminate misleading possibilities. Photographs of her Birkbeck work table [55]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. 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.”[56] However they knew they must complete their model before they could be certain.[57]

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.”[58] This was also one day after Franklin’s two A form papers had reached Acta Crystallogrphica. Wilkins came to see the model the following week, according to Maddox on 12 March, and allegedly informed Gosling on his return to King’s.[59]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,[60], 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.” [61]

Crick and Watson then published their model in Nature on 25 April 1953 in an article describing the double-helical structure of DNA with only a footnote acknowledging “having been stimulated by a general knowledge of” Franklin and Wilkin's ‘unpublished’ contribution.[62] 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 supported of the Crick and Watson theoretical paper which proposed a model for the B form of DNA.[63][64] Franklin left King's College London in March 1953 to move to Birkbeck College in a move that had been planned for some time.[43]

Weeks later, on 10 April, Franklin wrote to Crick for permission to see their model.[65] Franklin retained her scepticism for premature model building even after seeing the Crick-Watson 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 Crick-Watson model was in keeping with her cautious approach to science.[66]However, as documented above, she did not hesitate to publish preliminary ideas about DNA in ACTA, even before they could be definitively proven. Most of the scientific community hesitated several years before accepting the double helix proposal. At first mainly geneticists embraced the model because of its obvious genetic implications. Broader acceptance for the DNA double helix did not start until about 1960, and was not openly acknowledged until 1961 during the 1962 Nobel prize nominations. It took Wilkins and his colleagues approximately seven years to collect enough data to prove and refine the proposed DNA structure. According to the 1961 Crick-Monod letter cited above, 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.

[edit] Birkbeck College, London

Electronmicrograph of Tobacco Mosaic Virus

Franklin's work in Birkbeck involved the use of x-ray crystallography to study the structure of the tobacco mosaic virus (TMV) under J. D. Bernal[67] and was funded by the Agricultural Research Council (ARC).[68] In 1954 Franklin began a longstanding and successful collaboration with Aaron Klug.[69] In 1955 Franklin had a paper published in the journal Nature, indicating that TMV virus particles were all of the same length,[70] this was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct.[71]

Franklin worked on various viruses with different people. She eventually divided the work with her focusing on rod like viruses such as TMV (tobacco mosaic virus) with her PhD student Kenneth Holmes, while Aaron Klug worked on spherical viruses with his student John Finch, with Franklin coordinating and overseeing the work.[72] 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.[73] By the end of 1955 her team had completed a model of the TMV, to be exhibited at the upcoming Brussels World’s fair. The Birkbeck team members were working on viruses affecting several plants, including potato, turnip, tomato and pea.[74] Franklin and Don Caspar produced a paper each in Nature that taken together demonstrated that the RNA in TMV is wound along the inner surface of the hollow virus.[75][76]

[edit] Illness and death

In the summer of 1956, while on a work related trip to the USA, Franklin first began to suspect a health problem.[77] An operation in September of the same year revealed two tumours in her abdomen.[78] After this period and other periods of hospitalization, Franklin spent time convalescing with various friends and family. These included Anne Sayre, Crick and his wife Odile, with whom Franklin had formed a strong friendship,[79]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 a further six in 1957.[80] 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 USA.[81] At the end of 1957 Franklin again fell ill and was admitted to the Royal Marsden Hospital. She returned to work in January 1958 and was given a promotion to Research Associate in Biophysics.[82] She fell ill again on the 30th of March and died on 16 April 1958 in Chelsea, London,[83][84] of bronchopneumonia, secondary carcinomatosis and carcinoma of the ovary. Exposure to X-ray radiation is sometimes considered a possible factor in her illness. Other members of her family have died of cancer, and the incidence of "female" cancer is known to be disproportionately high amongst Ashkenazi Jews.[85] Her death certificate read: A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.[86]

[edit] Controversies after death

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

[edit] Sexism at King's College

Mainly because of mistaken impressions presented in Anne Sayre's 1975 Franklin biography, there have been assertions that Rosalind Franklin was discriminated against because of her gender and that King's, as an institution, was sexist. Among the examples cited in alleging sexist treatment at Kings was 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".[87][88] Actually these were the only options for a private lunch as most of the MRC group typically ate lunch together (including Franklin) in the mixed dining room discussed below[89]. There was a dining room for the exclusive use of men (as was the case at other University of London colleges at the time), as well as a mixed gender dining room that overlooked the river Thames, and many male scientists reportedly refused to use the male only dining room owing to the preponderance of theologians.[90]

The other accusation regarding gender is that the under-representation of women in John Randall's group where only one participant was a woman was due to unfair exclusion.[91] In contrast, defenders of the King's college MRC group correctly argue that women were (by the standards of the time) well-represented in the group, representing eight out of thirty-one members of staff,[92] or possibly closer to one in three,[93]although most were not senior scientists.[94]

[edit] Contribution to the model of DNA

Rosalind Franklin's critical contributions to the Crick and Watson model include an X-ray photograph of B-DNA (called photograph 51),[95] that was briefly shown to James Watson by Maurice Wilkins in January 1953,[96][97] and a report written for an MRC biophysics committee visit to King's in December 1952. This MRC report contained data from the King's group, including some of Rosalind Franklin's work, and was given to Francis Crick by his thesis supervisor Max Perutz, a member of the visiting committee.[98][99] Maurice Wilkins had been given photograph 51 by Rosalind Franklin's PhD student Raymond Gosling, because she was leaving King's to work at Birkbeck. There was nothing untoward in this transfer of data to Wilkins,[100][101]since Director Randall had insisted that all DNA work belonged exclusively to King's and had even instructed Franklin to stop thinking about it.[102]Also it was implied by Horace Freeland Judson, incorrectly, that Maurice Wilkins had taken the photograph out of Rosalind Franklin's drawer.[103] Likewise Max Perutz saw no harm in showing the MRC report to Crick as it had not been marked as confidential, although,"in the customary British manner in which everything official is considered secret until deliberately made public, the report was not expected to reach outside eyes."[104] 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 [105] and to post a general statement in "Science" excusing himself on the basis of being "inexperienced and casual in administrative matters." [106] Perutz also claimed that the MRC information was already made available to the Cambridge team when Watson had attended Franklin's seminar in November in 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 1951 November, which Watson had attended and not understood.[107][108] The Perutz letter was 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 1954 they were working with critical parameters determined by Franklin in 1951 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. Rosalind Franklin was probably never aware that her work had been used during construction of the model.[109]

[edit] Recognition of her contribution to the model of DNA

On the completion of their model, Francis Crick and James Watson had invited Maurice Wilkins to be a co-author of their paper describing the structure.[110][111] Wilkins turned down this offer, as he had taken no part in building the model.[112] Maurice Wilkins 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.[113] 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.[114]Indeed a clear timely acknowledgment would have been awkward, given the unorthodox manner in which data was 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 acknowledgment . "Such acknowledgment as they gave her was very muted and always coupled with the name of Wilkins."[115]

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 allegations that Franklin did not know how to interpret her own data and that she should have therefore shared her work with Wilkins, Watson and Crick. This attitude is epitomized in the confrontation between Watson and Franklin over a pre-print of Pauling's mistaken DNA manuscript.[116]Watson's words impelled Sayre to write her rebuttal, in which she designs her entire chapter nine, "Winner Take All" to be like a legal brief dissecting and analyzing the topic of acknowledgment.[117] Unfortunately Sayre’s early analysis was often ignored because of the supposed feminist overtones in her book. It should be noted that in their original paper, Watson and Crick do cite the the X-ray diffraction work of both Wilkins and William Astbury. In addition, they admit their ,"having been stimulated by a knowledge of the general nature of the unpublished experimental work of ..." groups led by both both Wilkins and Franklin.[2] Franklin and Raymond Gosling's own publication in the same issue of Nature was the first publication of this more clarified X-ray image of DNA.[118]

[edit] Nobel Prize

The rules of the Nobel Prize forbid posthumous nominations[111] and because Rosalind Franklin had died in 1958 she was not eligible for nomination to the Nobel Prize subsequently awarded to Crick, Watson, and Wilkins in 1962.[119] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.[120] By the time of the award Wilkins had been working on the structure of DNA for over 10 years, and had done much to confirm the Watson-Crick model.[121] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.[122]

[edit] Posthumous recognition

  • 2005, the wording on the DNA sculpture (which was donated by James Watson) outside Clare College's Thirkill Court, Cambridge, England 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." [1]
  • An Honorary 2008 Horwitz Prize will be awarded by Columbia University to Rosalind Franklin, Ph.D., posthumously, "for her seminal contributions to the discovery of the structure of DNA".[2]

[edit] Footnotes

  1. ^ Crick’s 31 December 1961 letter to Jacques Monod was discovered in the Archives of the Pasteur Institute by Doris Zeller, then reprinted in Nature Correspondence 425, 15 on the 4th of September 2003. Watson confirmed this opinion in his own statement at the opening of the King’s college Franklin-Wilkins building in 2000.
  2. ^ a b Watson JD, Crick FHC (1953). "A Structure for Deoxyribose Nucleic Acid". Nature 171: 737–738. Full text PDF This article was immediately followed by the two King’s submissions: M.H.F. Wilkins, A.R. Stokes, and H.R. Wilson. Molecular Structure of Deoxypentose Nucleic Acids , pp738-740 then by: Rosalind E. Franklin and R.G. Gosling. Molecular configuration of Sodium Thymonucleate pp 740-741.
  3. ^ Double Helix: 50 Years of DNA. Nature archives. Nature Publishing Group
  4. ^ Watson, James D. The Double Helix: A personal Account of the Discovery of the Structure of DNA (New York:Athenium,1968;London:Weidenfeldand Nicolson, 1981)
  5. ^ Consult references listed in order of publication: Olby, Sayre, Klug, Piper, Judson, Glynn, Maddox, Elkin
  6. ^ GRO Register of Births: SEP 1920 1a 250 KENSINGTON - Rosalind E. Franklin, mmn = Waley
  7. ^ Maddox, Brenda (2002). Rosalind Franklin: The Dark Lady of DNA. HarperCollins. ISBN 0060184078. 
  8. ^ Maddox p. 7
  9. ^ Segev p.
  10. ^ Sayre,A,. Rosalind Franklin and DNA(New York:Norton,1975). p. 31
  11. ^ a b Maddox p. 40
  12. ^ Maddox p. 25
  13. ^ Sayre p. 41
  14. ^ Maddox p. 30
  15. ^ Maddox, p. 26
  16. ^ Maddox, p. 20
  17. ^ Sayre, p. 35
  18. ^ Maddox, pp.40-82
  19. ^ Sayre pp. 47-57
  20. ^ Maddox, page 87
  21. ^ Maddox, p. 88
  22. ^ Maddox, p. 92
  23. ^ Franklin (1950)
  24. ^ Maddox, p. 108
  25. ^ Maddox, p. 111
  26. ^ Maddox, p. 124
  27. ^ Maddox, p. 114
  28. ^ Wilkins, Wilkins, M., The Third Man of the Double Helix, an autobiography (2003) Oxford University Press, Oxford. pp. 143-144
  29. ^ Wilkins, p. 121
  30. ^ 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).
  31. ^ Sayre, Olby, Maddox, Elkin, Wilkins
  32. ^ Maddox, p. 129
  33. ^ Elkin, p. 43
  34. ^ Wilkins p. 155
  35. ^ Elkin p. 45
  36. ^ a b Maddox, p. 153
  37. ^ Wilkins, p. 154
  38. ^ Maddox p 155
  39. ^ a b Wilkins, p. 158
  40. ^ Maddox, p. 155
  41. ^ a b Wilkins, p. 176
  42. ^ Wilkins, p. 182
  43. ^ a b Maddox, p. 168
  44. ^ Maddox, p. 169
  45. ^ Wilkins, pp. 232-233
  46. ^ Franklin, R.E. and Gosling, R.G. authors of papers received 6 March 1953 Acta Cryst. (1953). 6, 673 The Structure of Sodium Thymonucleate Fibres I. The Influence of Water Content Acta Cryst. (1953). 6, 678 The Structure of Sodium Thymonucleate Fibres II. The Cylindrically Symmetrical Patterson Function
  47. ^ Maddox p 205
  48. ^ Acta Cryst. (1953). 6, 673 The Structure of Sodium Thymonucleate Fibres I. The Influence of Water Content
  49. ^ Klug, A. “Rosalind Franklin and the Double Helix,” Nature 248 (26 April 1974): 787-788
  50. ^ Klug, A. Rosalind Franklin and the Discovery of the Structure of DNA, Nature 219 (24 August 1968): 808-810 & 843.
  51. ^ Yockey, pp. 9-10
  52. ^ Crick's 31 December 1961 letter to Jacque Monod cited above
  53. ^ Maddox, p. 147
  54. ^ Maddox, p. 161
  55. ^ Photograph was taken in the same series of photographs taken by Franklin’s colleague John Finch, as the one shown of Franklin’s Birkbeck desk that was reproduced by Maddox. Author Lynne Elkin was given a complete set and was informed by Finch that he had given a complete set to Maddox.
  56. ^ The Double Helix p. 115
  57. ^ The Double Helix p. 60
  58. ^ “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.
  59. ^ Maddox p. 207
  60. ^ In contrast to his other letters to Crick, Wilkins dated this one.
  61. ^ “Old rogues ” 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 208 and Olby.
  62. ^ Maddox, p. 212
  63. ^ Franklin and Gosling (1953)
  64. ^ Maddox, p. 210
  65. ^ 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.
  66. ^ Holt, J. (2002)
  67. ^ Maddox, p. 229
  68. ^ Maddox, p. 235
  69. ^ Maddox, p. 249
  70. ^ Franklin (1955)
  71. ^ Maddox, p. 252
  72. ^ Maddox, p. 254
  73. ^ Maddox, p. 256
  74. ^ Maddox, p. 262
  75. ^ Maddox, p. 269
  76. ^ Franklin (1956)
  77. ^ Maddox, p. 284
  78. ^ Maddox, p. 285
  79. ^ Maddox, p. 288
  80. ^ Maddox, p. 292
  81. ^ Maddox, p. 296
  82. ^ Maddox, p. 302
  83. ^ GRO Register of Deaths: JUN 1958 5c 257 CHELSEA - Rosalind E. Franklin, aged 37
  84. ^ Maddox, pp. 305-307
  85. ^ Maddox, p.320
  86. ^ Maddox, p.307
  87. ^ Sayre, p.97
  88. ^ Bryson, B. (2004) p. 490
  89. ^ Elkin, p.45
  90. ^ Maddox, p. 128
  91. ^ Sayre, p.99
  92. ^ Maddox, p. 133
  93. ^ Wilkins, p. 256
  94. ^ Elkin, p45
  95. ^ Maddox, pp. 177-178
  96. ^ Maddox, p. 196
  97. ^ Crick, (1988) p. 67.
  98. ^ Elkin, L.O. (2003)p 44
  99. ^ Maddox, pp. 198-199
  100. ^ Maddox, pp. 196
  101. ^ Wilkins, p. 198
  102. ^ Maddox p.312,
  103. ^ Wilkins, p. 257
  104. ^ Maddox p.188
  105. ^ Perutz papers are in the Archive of the J. Craig Venter institute and Science Foundation in Rockville Maryland, U.S.A., 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.
  106. ^ "Science, 27 June, 1969, pp. 207-212, also reprinted in the Norton critical edition of "The Double Helix," edited by Gunther Stent.
  107. ^ Maddox, p. 199
  108. ^ Watson (1969).
  109. ^ Maddox, p. 316
  110. ^ Wilkins, p. 213
  111. ^ a b Maddox, p. 205
  112. ^ Wilkins, p. 214
  113. ^ Wilkins, p. 226
  114. ^ Maddox, p. 207
  115. ^ Maddox, pp316-317, and other parts of the epilogue
  116. ^ Watson, J.D. (1968) pp. 95-96
  117. ^ Sayre,A. (1975) pp. 156-167
  118. ^ Franklin R, Gosling RG (1953) "Molecular Configuration in Sodium Thymonucleate". Nature 171: 740–741. Full text PDF
  119. ^ Nobel Prize (1962)
  120. ^ Wilkins, p. 242
  121. ^ Wilkins, p. 240
  122. ^ Wilkins, p. 243
  123. ^ Iota Sigma Pi professional awards recipients
  124. ^ a b c Maddox, p. 322
  125. ^ Sir Aaron Klug opens new Laboratory
  126. ^ NPG pictures
  127. ^ Maddox, p. 323
  128. ^ "seventh annual Rosalind E. Franklin Award for Women in Cancer Research at the National Cancer Institute's Intramural Scientific Retreat [which] honors the commitment of women in cancer research and is given in tribute to chemist Rosalind Franklin, who played a critical role in the discovery of the DNA double helix." The JHU Gazette, Johns Hopkins University, March 17 2008 For the Record: Cheers
  129. ^ The Royal Society Rosalind Franklin Award (2003): The Royal Society web page. Retrieved 21 July 2006.
  130. ^ Dedication of Rosalind Franklin University

[edit] References

  • Bryson, B. A Short History of Nearly Everything. (2004). Black Swan ISBN 0-552-99704-8.
  • Crick, F., H., C. and Watson, J., D. Molecular structure of nucleic acids (1953) Nature 171 pp. 737–738.
  • Crick, F. H. C. What Mad Pursuit, (1988). Basic Books. ISBN 0-465-09137-7.
  • Elkin, L., O. Rosalind Franklin and the Double Helix Physics Today March 2003, pp. 42–48.
  • Franklin RE (January 1950). "Influence of the bonding electrons on the scattering of X-rays by carbon". Nature 165 (4185): 71. PMID 15403103. 
  • 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.
  • Franklin, R and Gosling, R., G. Molecular Configuration in Sodium Thymonucleate Nature volume 171 pages 740-741. (1953)full text.
  • Franklin, R.E. and Gosling, R.G. authors of papers received 6 March 1953: Acta Cryst. (1953). 6, 673 The Structure of Sodium Thymonucleate Fibres I. The Influence of Water Content II. The Cylindrically Symmetrical Patterson Function
  • Franklin, R.E. "Structure of tobacco mosaic virus." Nature 175:379-381 (1955).
  • Franklin, R.E. (1956) "Location of the ribonucleic acid in the tobacco mosaic virus particle." Nature 177:928.
  • Holt, J. (2002) "Photo Finish: Rosalind Franklin and the great DNA race" The New Yorker October
  • Judson, Horace Freeland, "The Eighth Day of Creation:Makers of the Revolution in Biology" ( London: Jonathan Cape,1979),Penguin,1995;expanded edition;New York:Cold Spring Harbor Press,1996).
  • Maddox, B. Rosalind Franklin: The Dark Lady of DNA (2002). Harper Collins ISBN 0-00-655211-0.
  • Nobel Prize (1962). The Nobel Prize in Physiology or Medicine 1962, for their discoveries concerning the molecular structure of nucleic acids and its significance for information transfer in living material,
  • Olby, R., "The Path to the Double Helix"(London:Macmillan,1974)
  • Sayre, A. 1975. Rosalind Franklin and DNA. New York: W.W. Norton and Company. ISBN 0-393-32044-8.
  • Segev, T. One Palestine, Complete, (2000) (ISBN 0-349-11286-X) Abacus History.
  • Stent, Gunther, editor. "Critical Edition of The Double Helix"(1980) W.W. Norton Co, New York and London. ISBN 0-393-95075-1.
  • Watson, J. Letter to Science, 164, p. 1539, 27 (1969).
  • Wilkins, M., The Third Man of the Double Helix, an autobiography (2003) Oxford University Press, Oxford. ISBN 0-19-280667-X.
  • Yockey, H. P. Information Theory, Evolution, and the Origin of Life (2005).

[edit] Further reading

  • Brown, Andrew; "J. D. Bernal: The Sage of Science", Oxford University Press, 2005; ISBN 0-199-20565-5
  • Chomet, S. (Ed.), D.N.A. Genesis of a Discovery. Newman-Hemisphere Press (1994): NB a few copies are available from Newman-Hemisphere at 101 Swan Court, London SW3 5RY (phone/fax: 07092 060530).
  • Crick, Francis (1988) "What Mad Pursuit: A Personal View of Scientific Discovery" (Basic Books reprint edition, 1990) ISBN 0-465-09138-5
  • Dickerson, Richard E.; "Present at the Flood: How Structural Molecular Biology Came About", Sinauer, 2005; ISBN 0-878-93168-6;
  • Hager, Thomas; "Force of Nature: The Life of Linus Pauling", Simon & Schuster 1995; ISBN 0-684-80909-5
  • Freeland Judson, Horace (1996) [1977]. The Eighth Day of Creation: Makers of the Revolution in Biology (Expanded edition ed.). Plainview, N.Y: CSHL Press. ISBN 0-87969-478-5. 
  • Glynn, Jennifer Franklin. "Rosalind Franklin, 1920 - 1958" in "Cambridge Women: Twelve Portraits" (CUP 1996) pp 267 – 282 eds. Edward Shils and Carmen Blacker, ISBN 0521482879
  • Klug, A. Oxford Dictionary of National Biography article on R.E. Franklin, OUP, Matthew H.C.G. Ed., first published Sept 2004; online edn, Jan 2007, 1840 words; ISBN 019861411X; was selected "Life of The Day" on 16 April 2008 (50th anniversary of her death).
  • Klug, A. A lecture about Rosalind Franklin's contribution to the elucidation of the structure of DNA. in DNA Changing Science and Society: The Darwin Lectures for 2003 Krude, Torsten (Ed.) CUP (2003)
  • Olby, Robert, (1972) 'Rosalind Elsie Franklin' biography in "Dictionary of Scientific Biography", ed. Charles C. Gillespie (New York: Charles Scribner's sons) ISBN: ISBN 0684101211
  • Olby, Robert, The Path to The Double Helix: Discovery of DNA, (1974). MacMillan ISBN 0-486-68117-3
  • "Quiet debut for the double helix" by Professor Robert Olby, Nature 421 (January 23, 2003): 402-405.
  • Tait, Sylvia & James "A Quartet of Unlikely Discoveries" (Athena Press 2004) ISBN 184401343X
  • Watson, James D. (1980). The double helix: A personal account of the discovery of the structure of DNA. Norton. ISBN 0-393-01245-X. 
  • Wilkins, Maurice, "The Third Man of The Double Helix", OUP 2003; ISBN 978-0-19-280667-3.

[edit] External links

[edit] Articles

Franklin, S.*My aunt, the DNA pioneer

Piper, A.*Light on a Dark Lady, republished article from Trends in Biochemical Science

[edit] Documentaries

BBC docudrama (that is very accurate) "Life Story" aka in the USA "Race for the Double Helix"

[edit] Collections and publications

DNA structure research at King's College London 1947-1959
Rosalind Franklin | Raymond Gosling | John Randall | Alex Stokes | Maurice Wilkins | Herbert Wilson
NAME Franklin, Rosalind
ALTERNATIVE NAMES Franklin, Rosalind Elsie
SHORT DESCRIPTION biophysicist and crystallographer
DATE OF BIRTH 1920-07-25
PLACE OF BIRTH Notting Hill, London
DATE OF DEATH 1958-04-16
PLACE OF DEATH Chelsea, London

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