Rosalind Franklin

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Rosalind Franklin

Discovery of the DNA Double Helix

A recent painting of Franklin
Francis Crick
Rosalind Franklin
James Watson
Maurice Wilkins

Rosalind Elsie Franklin (25 July 1920 – 16 April 1958) was a British physical chemist and crystallographer who made important contributions to the understanding of the fine structures of DNA, viruses, coal and graphite. Franklin is best known for her contribution to the discovery of the structure of DNA in 1953. In the years following, she led pioneering work on the tobacco mosaic and polio viruses. She died in 1958 of cancer of the ovary.

Contents

[edit] Background

Franklin was born in London into an affluent and influential British-Jewish family.[1] Her great uncle was Herbert Samuel (later Viscount Samuel) who was Home Secretary in 1916 and the first practicing Jew to serve in the British Cabinet.[2] He was also the first High Commissioner (effectively governor) for the British Mandate of Palestine.

Her aunt Helen was married to Norman Bentwich who was Attorney General in the British Mandate of Palestine.[3] She was active in trade union organization and women's suffrage, and was later a member of the London County Council.[4][5]

Rosalind's uncle Hugh Franklin was also a supporter of women's suffrage, and spent six weeks in prison in 1910 for attacking Winston Churchill with a dog whip because of Churchill's opposition to this cause.[6][4]

Rosalind Franklin was educated at St Paul's Girls' School[7][8] where she excelled in Latin[9] and sport.[10] 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.[11][12] Later they helped settle Jewish refugees from Europe who had escaped the Nazis.[5]

[edit] Education and Career

[edit] University education

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

[edit] British Coal Utilisation Research Association

She worked for Ronald Norish between 1941 and 1942. Because of her desire to do war work during World War II, 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 doctoral degree-"The physical chemistry of solid organic colloids with special reference to coal and related materials" that she earned in 1945.[13][14]

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

After the war ended Franklin accepted an offer to work in Paris with Jacques Mering.[15] She learned x-ray diffraction techniques during her three years at the Laboratoire central des services chimiques de l'État.[16] She seemed to have been very happy there [17] and earned an international reputation based on her published research about the structure of coal.[18] In 1950 she sought work in England[19] and in June 1950 she was appointed to a position at King's College London.[20]

[edit] King's College London

In January 1951, Franklin started working as a research associate at King's College London in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall.[21] Although originally she was to have worked on x-ray diffraction of proteins in solution, her work was redirected to DNA fibers before she started working at King's.[22][23] Maurice Wilkins and Raymond Gosling had been carrying out x-ray diffraction analysis of DNA in the Unit since 1950. [24]
DNA pioneers
William Astbury
Oswald Avery
Erwin Chargaff
Max Delbrück
Jerry Donohue
Raymond Gosling
Phoebus Levene
Linus Pauling
Sir John Randall
Erwin Schrödinger
Alec Stokes
Herbert Wilson

Franklin, working with her student Raymond Gosling [25] started to apply the expertise she had gained in x-ray diffraction techniques to the structure of DNA. They 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.[26][27] These were termed DNA 'B' and 'A' respectively. The work on DNA was subsequently divided, Franklin taking the A form to study and Wilkins the 'B' form.[28][29] 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".[30]

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

By the end of 1951 it was generally accepted in King's that the B form of DNA was a helix, but Franklin in particular was unconvinced that the A form of DNA was helical in structure.[31] As a practical joke Franklin and Gosling produced a death notice regretting the loss of helical crystalline DNA (A-DNA).[32] During 1952 Rosalind Franklin and Raymond Gosling worked at applying the Patterson function to the x-ray pictures of DNA they had produced,[33] this was a long and labour-intensive approach but would give an insight into the structure of the molecule.[34][35]

In February 1953 Francis Crick and James Watson of the Cavendish Laboratory in Cambridge University had started to build a model of the B form of DNA using similar data to that available to the team at King's. Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,[36][37] but Rosalind Franklin was opposed to building theoretical models, taking the view that building a model was only to be undertaken after the structure was known.[38][39]
Watson and Crick then indirectly obtained a prepublication version of Franklin's DNA X-ray diffraction data, and a prepublication manuscript by Pauling and Corey, giving them critical insights into the DNA structure.[40]

Francis Crick and James Watson then published their model in Nature on 25 April 1953 in an article describing the double-helical structure of DNA with a small footnote to Franklin's data.[41] Articles by Maurice Wilkins and Rosalind Franklin [42] illuminating their x-ray diffraction data published in the same issue of Nature supported the Crick and Watson model for the B form of DNA.[43] Rosalind Franklin eventually left King's College London in March 1953 to move to Birkbeck College in a move that had been planned for some time.[44]

[edit] Birkbeck College, London

Electronmicrograph of Tobacco Mosaic Virus
Enlarge
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[45] and was funded by the Agricultural Research Council(ARC).[46] In 1954 Franklin began a longstanding and successful collaboration with Aaron Klug.[47] In 1955 Franklin had a paper published in the journal Nature, indicating that TMV virus particles were all of the same length, [48] this was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct.[49]

Franklin worked on rod like viruses such as TMV with her Ph.D. student Kenneth Holmes, while Aaron Klug worked on spherical viruses with his student John Finch, Franklin coordinated the work and was in charge.[50] 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.[51] By the end of 1955 her team had completed a model of the TMV and were working on viruses affecting several plants, including potato, turnip, tomato and pea.[52] Franklin and Don Casper 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.[53] [54]

[edit] Illness and death

In the summer of 1956, while on a work related trip to the United States of America (USA) Franklin first began to suspect a health problem.[55] An operation in September of the same year revealed two tumours in her abdomen.[56] After this period of illness Franklin spent some time convalescing at the home of Crick and his wife Odile.[57] She continued to work and her group continued to produce results, seven papers in 1956 and a further six in 1957.[58] 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.[59] 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.[60] She fell ill again on the 30th of March [61] and died on the 16th of April of cancer of the ovary.[62] Exposure to x-ray radiation is sometimes considered a possible factor in her illness, though she was no more reckless than other laboratory staff of the time. Other members of her family have died of cancer, and the incidence of cancer is known to be disproportionately high amongst Ashkenazi Jews.[63]

[edit] Controversies after death

Various controversies have surrounded Rosalind Franklin; these have all come to light after her death.

[edit] Allegations of sexism at King's College

There have been allegations that Rosalind Franklin was discriminated against because of her gender:

One of the allegations is that King's, as an institution, was sexist, the thrust of the allegation being that women were excluded from the staff dining room, and had to eat their meals in the student hall or away from the University. [64][65] This does not appear to be the whole story. Whereas it is true that there was a dining room for the exclusive use of men (as was the case at other University of London colleges at the time), there was also a mixed gender dining room that overlooked the river Thames, and many male scientists refused to use the male only dining room, due to the preponderance of theologians.[66]

The other accusation regarding gender is that women were under-represented in John Randall's group, the claim is that there was only one other woman in the group and that the women were excluded because of their gender.[67] In fact women seem to have been (by the standards of the time) exceptionally well-represented in the group, representing eight out of thirty-one members of staff,[68] or possibly closer to one in three.[69]

[edit] Contribution to the model of DNA

Rosalind Franklin's contributions to the Crick and Watson model include an X-ray photograph of B-DNA (called photograph 51),[70] that was briefly shown to James Watson by Maurice Wilkins in January 1953,[71][72] and a report written for an MRC biophysics committee visit to King's in December 1952. The 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.[73][74] 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,[75][76] though it has been implied, incorrectly, that Maurice Wilkins had taken the photograph out of Rosalind Franklin's drawer.[77] Likewise Max Perutz saw no harm in showing the MRC report to Francis Crick as it had not been marked as confidential. Much of the important material contained in the report had been presented by Franklin in a talk she had given in November 1951, which James Watson had attended.[78][79] The upshot of all this was that when Francis Crick and James Watson started to build their model in February 1953 they were working with 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.[80]

[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.[81][82] Maurice Wilkins turned down this offer, as he had taken no part in building the model.[83] Maurice Wilkins later expressed regret that greater discussion of co-authorship had not taken place as this may have helped to clarify the contribution the work at King's had made to the discovery.[84] There is no doubt that Rosalind Franklin's experimental data were used by Crick and Watson to build their model of DNA in 1953 (see above). That she is not cited in their original paper outlining their model may be a question of circumstance, it would have been very difficult to cite the unpublished work from the MRC report they had seen.[85] It should be noted that the x-ray diffraction work of both Maurice Wilkins and William Astbury are cited in the paper, and that the unpublished work of both Rosalind Franklin and Maurice Wilkins are acknowledged in the paper.[86]

[edit] Nobel Prize

The rules of the Nobel Prize forbid posthumous nominations.[87] Since Rosalind Franklin had died in 1958 she was not eligible for the Nobel Prize awarded to Francis Crick, James Watson and Maurice Wilkins in 1962.[88] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.[89] By the time of the award Maurice Wilkins had been working on the structure of DNA for over ten years, and had done much to confirm the Crick-Watson model.[90] Francis Crick had been working on the genetic code at Cambridge and James Watson had worked on RNA for some years.[91]

[edit] Posthumous recognition


[edit] References

  1. ^ Rosalind Franklin: The Dark Lady of DNA, p. 3
  2. ^ Rosalind Franklin: The Dark Lady of DNA, p. 7
  3. ^ Segev, Tom (2000): One Palestine, Complete, (ISBN 0-349-11286-X) Abacus History.
  4. ^ a b Rosalind Franklin and DNA, p. 31
  5. ^ a b Rosalind Franklin: The Dark Lady of DNA, p. 40
  6. ^ Rosalind Franklin: The Dark Lady of DNA, p. 28
  7. ^ Rosalind Franklin: The Dark Lady of DNA, p. 25
  8. ^ Rosalind Franklin and DNA, p. 41
  9. ^ Rosalind Franklin: The Dark Lady of DNA, p. 30
  10. ^ Rosalind Franklin: The Dark Lady of DNA, p. 26
  11. ^ Rosalind Franklin: The Dark Lady of DNA, p. 20
  12. ^ Rosalind Franklin and DNA, p. 35
  13. ^ Rosalind Franklin: The Dark Lady of DNA, pp.40-82
  14. ^ Rosalind Franklin and DNA pp. 47-57
  15. ^ Rosalind Franklin: The Dark Lady of DNA, page 87
  16. ^ Rosalind Franklin: The Dark Lady of DNA, p. 88
  17. ^ Rosalind Franklin: The Dark Lady of DNA, p. 92
  18. ^ R. E. Franklin: "Influence of the bonding electrons on the scattering of X-rays by carbon", Nature (1950) volume 165 page 71 Entrez PubMed 15403103
  19. ^ Rosalind Franklin: The Dark Lady of DNA, p. 108
  20. ^ Rosalind Franklin: The Dark Lady of DNA, p. 111
  21. ^ Rosalind Franklin: The Dark Lady of DNA, p. 124
  22. ^ Rosalind Franklin: The Dark Lady of DNA, p. 114
  23. ^ The Third Man of the Double Helix, pp. 143-144
  24. ^ The Third Man of the Double Helix, p. 121
  25. ^ Rosalind Franklin: The Dark Lady of DNA, p. 129
  26. ^ Rosalind Franklin: The Dark Lady of DNA, p. 153
  27. ^ The Third Man of the Double Helix, p. 154
  28. ^ The Third Man of the Double Helix, p. 158
  29. ^ Rosalind Franklin: The Dark Lady of DNA, p. 155
  30. ^ Rosalind Franklin: The Dark Lady of DNA, p. 153
  31. ^ The Third Man of the Double Helix, p. 176
  32. ^ The Third Man of the Double Helix, p. 182
  33. ^ Rosalind Franklin: The Dark Lady of DNA, p. 168
  34. ^ Rosalind Franklin: The Dark Lady of DNA, p. 169
  35. ^ The Third Man of the Double Helix, pp. 232-233
  36. ^ Rosalind Franklin: The Dark Lady of DNA, p. 147
  37. ^ The Third Man of the Double Helix, p. 158
  38. ^ Rosalind Franklin: The Dark Lady of DNA, p. 161
  39. ^ The Third Man of the Double Helix, p. 176
  40. ^ Hubert P. Yockey (2005)Information Theory, Evolution, and the Origin of Life, p9,10.
  41. ^ Rosalind Franklin: The Dark Lady of DNA, p. 212
  42. ^ R. Franklin and R. G. Gosling:Molecular Configuration in Sodium Thymonucleate, Nature (1953) volume 171 pages 740-741.full text
  43. ^ Rosalind Franklin: The Dark Lady of DNA, p. 210
  44. ^ Rosalind Franklin: The Dark Lady of DNA, p. 168
  45. ^ Rosalind Franklin: The Dark Lady of DNA, p. 229
  46. ^ Rosalind Franklin: The Dark Lady of DNA, p. 235
  47. ^ Rosalind Franklin: The Dark Lady of DNA, p. 249
  48. ^ Franklin, R.E. (1955) Structure of tobacco mosaic virus. Nature 175:379-381
  49. ^ Rosalind Franklin: The Dark Lady of DNA, p. 252
  50. ^ Rosalind Franklin: The Dark Lady of DNA, p. 254
  51. ^ Rosalind Franklin: The Dark Lady of DNA, p. 256
  52. ^ Rosalind Franklin: The Dark Lady of DNA, p. 262
  53. ^ Rosalind Franklin: The Dark Lady of DNA, p. 269
  54. ^ Franklin, R.E. (1956) Location of the ribonucleic acid in the tobacco mosaic virusparticle. Nature 177:928
  55. ^ Rosalind Franklin: The Dark Lady of DNA, p. 284
  56. ^ Rosalind Franklin: The Dark Lady of DNA, p. 285
  57. ^ Rosalind Franklin: The Dark Lady of DNA, p. 288
  58. ^ Rosalind Franklin: The Dark Lady of DNA, p. 292
  59. ^ Rosalind Franklin: The Dark Lady of DNA, p. 296
  60. ^ Rosalind Franklin: The Dark Lady of DNA, p. 302
  61. ^ Rosalind Franklin: The Dark Lady of DNA, p.305
  62. ^ Rosalind Franklin: The Dark Lady of DNA, p.307
  63. ^ Rosalind Franklin: The Dark Lady of DNA, p.320
  64. ^ Rosalind Franklin and DNA, p.97
  65. ^ Bryson, B. A Short History of Nearly Everything. (2004), page 490. Black Swan edition 0 552 99704 8
  66. ^ Rosalind Franklin: The Dark Lady of DNA, p. 128
  67. ^ Rosalind Franklin and DNA, p.99
  68. ^ Rosalind Franklin: The Dark Lady of DNA, p. 133
  69. ^ The Third Man of the Double Helix, p. 256
  70. ^ Rosalind Franklin: The Dark Lady of DNA, pp. 177-178
  71. ^ Rosalind Franklin: The Dark Lady of DNA, p. 196
  72. ^ Crick, F. H. C. What Mad Pursuit, p67 ISBN 0-465-09137-7
  73. ^ Elkin, L.O. Rosalind Franklin and the Double Helix Physics Today March 2003, p. 61
  74. ^ Rosalind Franklin: The Dark Lady of DNA, pp. 198-199
  75. ^ Rosalind Franklin: The Dark Lady of DNA, pp. 196
  76. ^ The Third Man of the Double Helix, p. 198
  77. ^ The Third Man of the Double Helix, p. 257
  78. ^ Rosalind Franklin: The Dark Lady of DNA, p. 199
  79. ^ Watson, James: Letter to Science, 164, p. 1539, 27 June 1969.
  80. ^ Rosalind Franklin: The Dark Lady of DNA, p. 316
  81. ^ The Third Man of the Double Helix, p. 213
  82. ^ Rosalind Franklin: The Dark Lady of DNA, p. 205
  83. ^ The Third Man of the Double Helix, p. 214
  84. ^ The Third Man of the Double Helix, p. 226
  85. ^ Rosalind Franklin: The Dark Lady of DNA, p. 207
  86. ^ Crick, F. H. C. and Watson, J. D. (1953) Molecular structure of nucleic acids, Nature 171 pp. 737-738
  87. ^ Rosalind Franklin: The Dark Lady of DNA, p. 205
  88. ^ 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, Nobelprize.org
  89. ^ The Third Man of the Double Helix, p. 242
  90. ^ The Third Man of the Double Helix, p. 240
  91. ^ The Third Man of the Double Helix, p. 243
  92. ^ a b c Rosalind Franklin: The Dark Lady of DNA, p. 322
  93. ^ Sir Aaron Klug opens new Laboratory
  94. ^ NPG pictures
  95. ^ Rosalind Franklin: The Dark Lady of DNA, p. 323
  96. ^ The Royal Society Rosalind Franklin Award (2003): The Royal Society web page. Retrieved 21 July 2006.
  97. ^ Dedication of Rosalind Franklin University

[edit] Major sources

[edit] Further reading

  • Watson, James D. (1980). The double helix: A personal account of the discovery of the structure of DNA. Norton. ISBN 0-393-01245-X.
  • Chomet, S. (Ed.), D.N.A. Genesis of a Discovery, (1994). Newman-Hemisphere Press: www.pantadeusz.zoomshare.com or e-mail: publications@zoomshare.com or phone/fax: 07092 060530
  • Krude, Torsten (Ed.) DNA Changing Science and Society: The Darwin Lectures for 2003 CUP 2003, includes a lecture by Sir Aaron Klug on Rosalind Franklin's involvement in the determination of the structure of DNA.
  • Freeland Judson, Horace, The Eighth Day of Creation: Makers of the Revolution in Biology (1995) Penguin Books. First published (1977) Jonathan Cape. ISBN 13579108642.

[edit] See also

[edit] External links

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