Rosalind Franklin

Rosalind Elsie Franklin (25 July 1920 – 16 April 1958) was a British biophysicist, physicist, chemist, biologist and X-ray crystallographer who made contributions to the understanding of the fine molecular structures of DNA, RNA, viruses, coal and graphite. Franklin is best known for her work on the X-ray diffraction images of DNA. Her data, according to Francis Crick, was a part of the data used to formulate Crick and Watson's 1953 hypothesis regarding the structure of DNA.^[1][2] Unpublished drafts of her papers (written as she was arranging to leave King's College London) show that she had determined the overall B-form of the DNA helix. Her work supported the hypothesis of Watson and Crick and was published third, in the series of three DNA Nature articles.^[3] 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^[4] into an affluent and influential British-Jewish family.^[5] 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 from Herbert Samuel (later Viscount Samuel) who was Home Secretary in 1916 and the first practising Jew to serve in the British Cabinet. He was also the first High Commissioner (effectively governor) for the British Mandate of Palestine.^[6]

Her aunt Helen Carolin Franklin was married to Norman Bentwich, who was Attorney General in the British Mandate of Palestine. She was active in trade union organisation and women's suffrage, and was later a member of the London County Council.^[7][8][9]

Franklin was educated at St Paul's Girls' School, where she excelled in science, Latin, and sports. 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. Later Franklin's family helped settle Jewish refugees from Europe who had escaped the Nazis.^{[9][10][11][12][13]}

In the winter 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 Ohio 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." Working with Jacques Mering, 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. She became known as an outstanding X-ray crystallographer, and was later recruited eagerly by the King's College group for their DNA project. Franklin seemed to have been very happy in France, and earned an international reputation based on her published research on the structure of coal. In 1950 she sought work in the UK and in June 1950 she was appointed to a position at King's College London.^[14][15][16]

[edit] King's College London

Double Helix Discovery
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

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

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.^[17] 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.^[18][19] He made this reassignment, even before she started working at King's, because of the following pioneering work by Maurice Wilkins and Raymond Gosling.^[20] 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.^[21] Randall’s lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin.^[22] Franklin, working with her student Raymond Gosling,^[23] 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. She manipulated the hydration of her specimens.^[24] When Wilkins inquired about this improved technique, Franklin replied in terms which offended Wilkins as Franklin had “an air of cool superiority.”^[25][26] 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.^[27][28] These were termed DNA 'B' and 'A' respectively. Because of the intense personality conflict developing between Franklin and Wilkins, Randall ^[29] divided the work on DNA. Franklin chose the data rich A form while Wilkins selected the 'B' form^[30][31] 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".^[27]

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^[32] 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).^[33] During 1952 Rosalind Franklin and Raymond Gosling worked at applying the Patterson function to the x-ray pictures of DNA they had produced.^[34] This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule.^[35][36]

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,^[37] one day before Crick and Watson had completed their model.^[38] 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.^[39] 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.^[40] Klug designed this paper to complement the first article he had written defending Franklin’s significant contribution to DNA structure.^[41]. 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. Wilkins 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.^[42] Franklin and Gosling’s photo 51 gave the Cambridge pair insights into the DNA structure, whereas Pauling and Corey’s paper described a molecule remarkably like their first incorrect model. The results of Franklin's constituted contributions to the Crick and Watson model. When Crick and Watson started to build their model in February 1954 they were working with parameters that had been determined by Franklin in 1951, and which she and Raymond Gosling had refined in 1952. The data included an X-ray photograph of B-DNA (called photograph 51), that was briefly shown to James Watson by Maurice Wilkins in January 1953, and an unpublished report written for an MRC biophysics committee visit to King's in December 1952 which was shown by Dr. Max Perutz at the Cavendish Laboratory to both Crick and Watson.^[43][44][45] This MRC report contained data from the King's group, including some of Rosalind Franklin's and Gosling's work, and was given to Francis Crick — who was working on his thesis on haemoglobin structure — by his thesis supervisor Max Perutz, a member of the visiting committee.^[46][47] Maurice Wilkins had been given photograph 51 by Rosalind Franklin's Ph.D. student Raymond Gosling per instructions of Director John Randall because Franklin was leaving King's to work at Birkbeck.^[48][49][50] 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 most complete set of data by February 1953.^[51] Model building had been applied successfully in the elucidation of the structure of the alpha helix by Linus Pauling in 1951,^[30][52] but Franklin was opposed to prematurely building theoretical models, until sufficient data was obtained properly to guide the model building. She took the view that building a model was to be undertaken only after enough of the structure was known.^[32][53] Ever cautious, she wanted to eliminate misleading possibilities. Photographs of her Birkbeck work table ^[54] 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.^[55] 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 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.^[34]

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.

Birkbeck College~ DNA Discoveries

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) as a senior scientist with her own research group, funded by the Agricultural Research Council (ARC). She was recruited by physics department chair J. D. Bernal, a brilliant crystallographer who happened to be an Irish communist, known for promoting women crystallographers. In 1954 Franklin began a longstanding and successful collaboration with Aaron Klug. In 1955 Franklin had a paper published in the journal Nature, indicating that TMV virus particles were all of the same length, this was in direct contradiction to the ideas of the eminent virologist Norman Pirie, though her observation ultimately proved correct. Franklin, and the research group she headed, focused on the structure of DNA. DNA actually constitutes the genome (central information molecule) of many viruses, including tobacco mosaic virus and polio virus. She assigned the study of rod-like viruses such as TMV (tobacco mosaic virus) to her PhD student Kenneth Holmes, while her colleague Aaron Klug worked on spherical viruses with his student John Finch, with Franklin coordinating and overseeing the work. 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. 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 DNA viruses affecting several plants, including potato, turnip, tomato and pea. Franklin and Don Caspar produced a paper each in Nature that taken together demonstrated that the DNA in TMV is wound along the inner surface of the hollow virus.^[67][68][69]

Her former colleagues at Birkbeck College, London Aaron Klug, John Finch and Kenneth Holmes moved to the Laboratory of Molecular Biology, Cambridge in 1962; had she lived, Rosalind Franklin would have moved with them, according to the scientific historian Robert Olby.

[edit] Illness and death

In the summer of 1956, while on a work-related trip to the United States, Franklin first began to suspect a health problem. An operation in September of the same year revealed two tumours in her abdomen. 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, 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. 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. At the end of 1957, Franklin again fell ill and she was admitted to the Royal Marsden Hospital. She returned to work in January 1958 and she was given a promotion to Research Associate in Biophysics. She fell ill again on March 30 and died on April 16, 1958, in Chelsea, London, 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 some sort of cancer, and the incidence of ovarian, uterine and breast cancers is known to be disproportionately high among Ashkenazi Jews. Her death certificate read: A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.^[70][71]

[edit] Controversies after death

Various controversies surrounding Rosalind Franklin were raised following her death.

[edit] Contribution to the model of DNA

Rosalind Franklin was probably never aware that her work had been used during construction of the model, but Maurice Wilkins was, and some feel that this was inappropriate.^[72] It was later implied incorrectly by Horace Freeland Judson that Maurice Wilkins had taken the photograph out of Rosalind Franklin's desk drawer, causing an uproar.^[73] Others have suggested that not obtaining Franklin's permission before sharing the results was somehow an oversight. It was later suggested by some that since a customary British notion is that everything official is considered secret until it is deliberately made public, that Perutz's sharing of the MRC report should have been treated as confidential even though it was not marked as such.^[74]

After the publication of Watson's The Double Helix, Perutz was beset by so many letters questioning his judgement that he felt the need to both answer them all and to post a general statement in Science excusing himself on the basis of being "inexperienced and casual in administrative matters." ^[75][76] Perutz also indicated that the MRC information was already made available to the Cambridge team when Watson had attended Franklin's seminar in November 1951.^[77] 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.^[78]

[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 but he turned down this offer, as he had taken no part in conceiving of the model.^[79][80] 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, but a full citation was not possible since Franklin's experimental data in the MRC report remained unpublished.^[81][82] Watson and Crick did not cite the MRC report as a personal communication or else cited the ACTA articles in press nor did they cite the third Nature paper that they knew was in press. What acknowledgement Franklin did receive was always coupled with the name of Wilkins.^[83]

In Watson's account, The Double Helix, he gave a clear recitation of Franklin's contribution. Watson asserted 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. He also mentions a disagreement about acknowledgement between himself and Franklin over a pre-print of Pauling's mistaken DNA manuscript.^[84][85] In their original paper, Watson and Crick do cite 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.^[86]

[edit] Nobel Prize

The rules of the Nobel Prize forbid posthumous nominations^[80] 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.^[87] The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA.^[88] 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.^[89] Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.^[90]

[edit] As a victim of sexism

While Franklin never complained of sexism during her lifetime, others have claimed that she was a victim of sexism. Some cite The Double Helix, in which Watson denigrates her work and frequently refers to her in patronizing terms as "Rosie," a name she never used. Much later, Francis Crick acknowledges, "I'm afraid we always used to adopt--let's say, a patronizing attitude towards her." Cambridge colleague Peter Cavendish wrote in a letter, "Wilkins is supposed to be doing this work; Miss Franklin is evidently a fool." The one laboratory director who supposedly supported her, John Randall, pointedly told her to "cease to work on the nucleic acid problem" upon her departure from King's.^[5]

Some have claimed that King's College was the source of the sexism. Their primary evidence is a male-only large comfortable dining room (rather common at other University of London colleges at the time) used by the preponderance of theologians while there was also supplied a mixed-gender dining room that overlooked the River Thames was the more typical lunch venue for the MRC group (and Franklin) and other male scientists. The female staff of all ranks "lunched in the students' hall or away from the premises".^{[91][92][93][94]} Others have cited the fact that John Randall's group had only one woman.^[95] while overall the King's college MRC group and staff had eight out of thirty-one female members, of which most were not senior scientists.^[96][97][98]

[edit] Posthumous recognition

Rosalind Franklin University of Medicine and Science

• 1982, Iota Sigma Pi designated Franklin a National Honorary Member.^[99]
• 1992, English Heritage placed a blue plaque on the house Rosalind Franklin grew up in.^[100]
• 1993, King's College London rename the Orchard Residence at their Hampstead Campus on Kidderpore Avenue Rosalind Franklin Hall.
• 1995, Newnham College dedicated a residence in her name and put a bust of her in its garden.^[100]
• 1997, Birkbeck, University of London School of Crystallography opened the Rosalind Franklin laboratory.^[101]
• 1998, National Portrait Gallery added Rosalind Franklin's next to those of Francis Crick, James Watson and Maurice Wilkins.^[100][102]
• 2000, King's College London opened the Franklin-Wilkins Building in honour of Dr. Franklin's and Professor Wilkins's work at the college.^[103] King's had earlier, in 1994, also named one of the Halls in Hampstead Campus residences in memory of Rosalind Franklin.
• 2001, The U.S. National Cancer Institute established the Rosalind E. Franklin Award for Women in Science.^[104]
• 2003, the Royal Society established the Rosalind Franklin Award, for an outstanding contribution to any area of natural science, engineering or technology.^[105]
• 2004, Finch University of Health Sciences/The Chicago Medical School, located in North Chicago, IL, changed its name to Rosalind Franklin University of Medicine and Science.^[106]
• 2004, The University of Groningen, The Netherlands installed Rosalind Franklin fellowships to promote the hiring of young, promising, female researchers.
• 2005, the wording on the DNA sculpture (which was donated by James Watson) outside Clare College's Thirkill Court, Cambridge, UK 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."^[107]
• 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".^[108]

[edit] List of publications

• 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.

[edit] Footnotes

[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.
• 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, Nobelprize.org
• 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;
• John Finch; 'A Nobel Fellow On Every Floor', Medical Research Council 2008, 381 pp, ISBN 978-1840469-40-0; this book is all about the MRC Laboratory of Molecular Biology, Cambridge
• 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

Wikimedia Commons has media related to: Rosalind Franklin

• Rosalind Franklin in CWP at UCLA
• Himetop – The History of medicine topographical database in http://himetop.wikidot.com/rosalind-franklin
• The People's Archive: Video interview with Aaron Klug: video segments relating to Rosalind Franklin:
  • 12. Work at Birbeck and meeting Franklin
  • 13. Work with Franklin
  • 17. Franklin and the discovery of the structure of DNA
  • 18. Franklin's death and joining the MRC's LMB in Cambridge

[edit] Articles

Franklin, S.*My aunt, the DNA pioneer

• Elkin, L.O. "Rosalind Franklin and the Double Helix" in "Physics Today" magazine

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

• Oxford Dictionary of National Biography for article by Sir Aaron Klug
• Clue to chemistry of heredity found The New York Times, 1953-06-13

[edit] Documentaries

• Nova: "Secret of Photo 51"

[edit] Collections and publications

• The Rosalind Franklin Society.
• The Rosalind Franklin Papers at the National Library of Medicine
• Anne Sayre Collection of Rosalind Franklin Materials
• Lynne Elkin collection of original Franklin research at the Bancroft Archive of the University of California, Berkeley
• The Rosalind Franklin Papers at Churchill Archives Centre in Cambridge.
• 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 | Alex Stokes | Maurice Wilkins | Herbert Wilson