Rosalind Franklin
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Rosalind Elsie Franklin (25 July 192016 April 1958) was a British
chemist A chemist (from Greek ''chēm(ía)'' alchemy; replacing ''chymist'' from Medieval Latin ''alchemist'') is a scientist trained in the study of chemistry. Chemists study the composition of matter and its properties. Chemists carefully describe th ...
and
X-ray crystallographer X-ray crystallography is the experimental science determining the atomic and molecular structure of a crystal, in which the crystalline structure causes a beam of incident X-rays to diffract into many specific directions. By measuring the angle ...
whose work was central to the understanding of the molecular structures of DNA (deoxyribonucleic acid), RNA (ribonucleic acid), viruses, coal, and
graphite Graphite () is a crystalline form of the element carbon. It consists of stacked layers of graphene. Graphite occurs naturally and is the most stable form of carbon under standard conditions. Synthetic and natural graphite are consumed on la ...
. Although her works on coal and viruses were appreciated in her lifetime, her contributions to the discovery of the structure of DNA were largely unrecognized during her life, for which she has been variously referred to as the "wronged heroine", the "dark lady of DNA", the "forgotten heroine", a "feminist icon", and the " Sylvia Plath of molecular biology". She graduated in 1941 with a degree in
natural sciences Natural science is one of the branches of science concerned with the description, understanding and prediction of natural phenomena, based on empirical evidence from observation and experimentation. Mechanisms such as peer review and repeat ...
from
Newnham College, Cambridge Newnham College is a women's constituent college of the University of Cambridge. The college was founded in 1871 by a group organising Lectures for Ladies, members of which included philosopher Henry Sidgwick and suffragist campaigner Millic ...
, and then enrolled for a PhD in
physical chemistry Physical chemistry is the study of macroscopic and microscopic phenomena in chemical systems in terms of the principles, practices, and concepts of physics such as motion, energy, force, time, thermodynamics, quantum chemistry, statistica ...
under Ronald George Wreyford Norrish, the 1920 Chair of Physical Chemistry at the
University of Cambridge , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
. Disappointed by Norrish's lack of enthusiasm, she took up a research position under the
British Coal Utilisation Research Association British Coal Utilisation Research Association (BCURA) was a non-profit association of industrial companies, incorporated 23 April 1938 and dissolved 24 February 2015. History It was founded in 1938, with an assured income of £25000 per year for ...
(BCURA) in 1942. The research on coal helped her earn a PhD from Cambridge in 1945. Moving to Paris in 1947 as a (postdoctoral researcher) under Jacques Mering at the ''Laboratoire Central des Services Chimiques de l'État'', she became an accomplished X-ray crystallographer. After joining
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
in 1951 as a research associate, she discovered the key properties of DNA, which eventually facilitated the correct description of the double helix structure of DNA. Owing to disagreement with her director, John Randall, and her colleague Maurice Wilkins, she was compelled to move to
Birkbeck College , mottoeng = Advice comes over nightTranslation used by Birkbeck. , established = , type = Public research university , endowment = £4.3 m (2014) , budget = £10 ...
in 1953. Franklin is best known for her work on the X-ray diffraction images of DNA while at King's College London, particularly Photo 51, taken by her student Raymond Gosling, which led to the discovery of the DNA double helix for which
Francis Crick Francis Harry Compton Crick (8 June 1916 – 28 July 2004) was an English molecular biologist, biophysicist, and neuroscientist. He, James Watson, Rosalind Franklin, and Maurice Wilkins played crucial roles in deciphering the helical stru ...
,
James Watson James Dewey Watson (born April 6, 1928) is an American molecular biologist, geneticist, and zoologist. In 1953, he co-authored with Francis Crick the academic paper proposing the double helix structure of the DNA molecule. Watson, Crick a ...
, and Maurice Wilkins shared the
Nobel Prize in Physiology or Medicine The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accordi ...
in 1962. Watson suggested that Franklin would have ideally been awarded a
Nobel Prize in Chemistry ) , image = Nobel Prize.png , alt = A golden medallion with an embossed image of a bearded man facing left in profile. To the left of the man is the text "ALFR•" then "NOBEL", and on the right, the text (smaller) "NAT•" then "M ...
, along with Wilkins but, although there was not yet a rule against posthumous awards, the Nobel Committee generally did not make posthumous nominations. Working under John Desmond Bernal, Franklin led pioneering work at Birkbeck on the molecular structures of viruses. On the day before she was to unveil the structure of tobacco mosaic virus at an international fair in Brussels, she died of
ovarian cancer Ovarian cancer is a cancerous tumor of an ovary. It may originate from the ovary itself or more commonly from communicating nearby structures such as fallopian tubes or the inner lining of the abdomen. The ovary is made up of three different ...
at the age of 37 in 1958. Her team member
Aaron Klug Sir Aaron Klug (11 August 1926 – 20 November 2018) was a British biophysicist and chemist. He was a winner of the 1982 Nobel Prize in Chemistry for his development of crystallographic electron microscopy and his structural elucidation of bi ...
continued her research, winning the Nobel Prize in Chemistry in 1982.


Early life

Franklin was born on 25 July 1920 in 50 Chepstow Villas, Notting Hill, London, into an affluent and influential
British Jewish British Jews (often referred to collectively as British Jewry or Anglo-Jewry) are British citizens who identify as Jewish. The number of people who identified as Jews in the United Kingdom rose by just under 4% between 2001 and 2021. History ...
family.


Family

Franklin's father was Ellis Arthur Franklin (1894–1964), a politically liberal London merchant banker who taught at the city's Working Men's College, and her mother was Muriel Frances Waley (1894–1976). Rosalind was the elder daughter and the second child in the family of five children. David (1919–1986) was the eldest brother; Colin (1923–2020),
Roland Roland (; frk, *Hrōþiland; lat-med, Hruodlandus or ''Rotholandus''; it, Orlando or ''Rolando''; died 15 August 778) was a Frankish military leader under Charlemagne who became one of the principal figures in the literary cycle known as the ...
(born 1926), and Jenifer (born 1929) were her younger siblings.Glynn, p. 1. Franklin's paternal great-uncle was Herbert Samuel (later Viscount Samuel), who was the
Home Secretary The secretary of state for the Home Department, otherwise known as the home secretary, is a senior minister of the Crown in the Government of the United Kingdom. The home secretary leads the Home Office, and is responsible for all nationa ...
in 1916 and the first practising Jew to serve in the British Cabinet.Maddox, p. 7. Her aunt, Helen Caroline Franklin, known in the family as Mamie, was married to
Norman de Mattos Bentwich Norman de Mattos Bentwich (28 February 1883 – 8 April 1971) was a British barrister and legal academic. He was the British-appointed attorney-general of Mandatory Palestine and a lifelong Zionist. Biography Early life Norman Bentwich was ...
, who was the
Attorney General In most common law jurisdictions, the attorney general or attorney-general (sometimes abbreviated AG or Atty.-Gen) is the main legal advisor to the government. The plural is attorneys general. In some jurisdictions, attorneys general also have exec ...
in the British Mandate of Palestine. Helen was active in trade union organisation and the
women's suffrage movement Women's suffrage is the right of women to vote in elections. Beginning in the start of the 18th century, some people sought to change voting laws to allow women to vote. Liberal political parties would go on to grant women the right to vot ...
and was later a member of the
London County Council London County Council (LCC) was the principal local government body for the County of London throughout its existence from 1889 to 1965, and the first London-wide general municipal authority to be directly elected. It covered the area today kn ...
.Maddox, p. 40. Franklin's uncle, Hugh Franklin, was another prominent figure in the suffrage movement, although his actions therein embarrassed the Franklin family. Rosalind's middle name, "Elsie", was in memory of Hugh's first wife, who died in the 1918 flu pandemic. Her family was actively involved with the Working Men's College, where her father taught the subjects of electricity, magnetism, and the history of the Great War in the evenings, later becoming the vice principal. Franklin's parents helped settle Jewish refugees from Europe who had escaped the
Nazis Nazism ( ; german: Nazismus), the common name in English for National Socialism (german: Nationalsozialismus, ), is the far-right totalitarian political ideology and practices associated with Adolf Hitler and the Nazi Party (NSDAP) in N ...
, particularly those from the '' Kindertransport''. They took in two Jewish children to their home, and one of them, a nine-year-old Austrian, Evi Eisenstädter, shared Jenifer's room. (Evi's father Hans Mathias Eisenstädter had been imprisoned in Buchenwald, and after liberation, the family adopted the surname "Ellis".)


Education

From early childhood, Franklin showed exceptional scholastic abilities. At age six, she joined her brother Roland at Norland Place School, a private day school in West London. At that time, her aunt Mamie (Helen Bentwich), described her to her husband: "Rosalind is alarmingly clever – she spends all her time doing arithmetic for pleasure, and invariably gets her sums right." She also developed an early interest in
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by st ...
and hockey. At age nine, she entered a boarding school, Lindores School for Young Ladies in Sussex. The school was near the seaside, and the family wanted a good environment for her delicate health. She was 11 when she went to St Paul's Girls' School in
Hammersmith Hammersmith is a district of West London, England, southwest of Charing Cross. It is the administrative centre of the London Borough of Hammersmith and Fulham, and identified in the London Plan as one of 35 major centres in Greater London ...
, west London, one of the few girls' schools in London that taught physics and chemistry. At St Paul's she excelled in science, Latin, and sports. She also learned German, and became fluent in French, a language she would later find useful. She topped her classes, and won annual awards. Her only educational weakness was in music, for which the school music director, the composer Gustav Holst, once called upon her mother to enquire whether she might have suffered from hearing problems or
tonsillitis Tonsillitis is inflammation of the tonsils in the upper part of the throat. It can be acute or chronic. Acute tonsillitis typically has a rapid onset. Symptoms may include sore throat, fever, enlargement of the tonsils, trouble swallowing, a ...
. With six distinctions, she passed her matriculation in 1938, winning a scholarship for university, the School Leaving Exhibition of £30 a year for three years, and £5 from her grandfather. Her father asked her to give the scholarship to a deserving refugee student.


Cambridge and World War II

Franklin went to
Newnham College, Cambridge Newnham College is a women's constituent college of the University of Cambridge. The college was founded in 1871 by a group organising Lectures for Ladies, members of which included philosopher Henry Sidgwick and suffragist campaigner Millic ...
, in 1938 and studied chemistry within the
Natural Sciences Tripos The Natural Sciences Tripos (NST) is the framework within which most of the science at the University of Cambridge is taught. The tripos includes a wide range of Natural Sciences from physics, astronomy, and geoscience, to chemistry and biology, ...
. There she met the spectroscopist Bill Price, who worked with her as a laboratory demonstrator and who later became one of her senior colleagues at King's College London. In 1941, she was awarded
second-class honours The British undergraduate degree classification system is a grading structure for undergraduate degrees or bachelor's degrees and integrated master's degrees in the United Kingdom. The system has been applied (sometimes with significant variati ...
from her final exams. The distinction was accepted as a bachelor's degree in qualifications for employment. Cambridge began awarding titular BA and MA degrees to women from 1947, and the previous women graduates retroactively received these. In her last year at Cambridge, she met a French refugee Adrienne Weill, a former student of
Marie Curie Marie Salomea Skłodowska–Curie ( , , ; born Maria Salomea Skłodowska, ; 7 November 1867 – 4 July 1934) was a Polish and naturalized-French physicist and chemist who conducted pioneering research on radioactivity. She was the fir ...
, who had a huge influence on her life and career and who helped her to improve her conversational French. Franklin was awarded a research fellowship at Newnham College, with which she joined the physical chemistry laboratory of the
University of Cambridge , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
to work under Ronald George Wreyford Norrish, who later won the
Nobel Prize in Chemistry ) , image = Nobel Prize.png , alt = A golden medallion with an embossed image of a bearded man facing left in profile. To the left of the man is the text "ALFR•" then "NOBEL", and on the right, the text (smaller) "NAT•" then "M ...
. In her one year of work there, she did not have much success. As described by his biographer, Norrish was "obstinate and almost perverse in argument, overbearing and sensitive to criticism". He could not decide upon the assignment of work for her. At that time he was succumbing to heavy drinking. Franklin wrote that he made her despise him completely. Resigning from Norrish's Lab, she fulfilled the requirements of the National Service Acts by working as an assistant research officer at the
British Coal Utilisation Research Association British Coal Utilisation Research Association (BCURA) was a non-profit association of industrial companies, incorporated 23 April 1938 and dissolved 24 February 2015. History It was founded in 1938, with an assured income of £25000 per year for ...
(BCURA) in 1942. The BCURA was located on the Coombe Springs Estate near
Kingston upon Thames Kingston upon Thames (hyphenated until 1965, colloquially known as Kingston) is a town in the Royal Borough of Kingston upon Thames, southwest London, England. It is situated on the River Thames and southwest of Charing Cross. It is notable as ...
near the southwestern boundary of London. Norrish acted as advisor to the military at BCURA. John G. Bennett was the director.
Marcello Pirani Marcello Stefano Pirani (July 1, 1880 – January 11, 1968) was a German physicist known for his invention of the Pirani vacuum gauge, a vacuum gauge based on the principle of heat loss measurement. Throughout his career, he worked on advancing l ...
and
Victor Goldschmidt Victor Moritz Goldschmidt (27 January 1888 in Zürich – 20 March 1947 in Oslo) was a Norwegian mineralogist considered (together with Vladimir Vernadsky) to be the founder of modern geochemistry and crystal chemistry, developer of the Golds ...
, both refugees from the Nazis, were consultants and lectured at BCURA while Franklin worked there. During her BCURA research, she initially stayed at Adrienne Weill's boarding house in Cambridge until her cousin, Irene Franklin, proposed that they share living quarters at a vacated house in Putney that belonged to her uncle. With Irene, she volunteered as an
Air Raid Warden Air Raid Precautions (ARP) refers to a number of organisations and guidelines in the United Kingdom dedicated to the protection of civilians from the danger of air raids. Government consideration for air raid precautions increased in the 1920s an ...
and regularly made patrols to see the welfare of people during air raids. She studied the
porosity Porosity or void fraction is a measure of the void (i.e. "empty") spaces in a material, and is a fraction of the volume of voids over the total volume, between 0 and 1, or as a percentage between 0% and 100%. Strictly speaking, some tests measur ...
of coal using helium to determine its density. Through this, she discovered the relationship between the fine constrictions in the pores of coals and the permeability of the porous space. By concluding that substances were expelled in order of molecular size as temperature increased, she helped classify coals and accurately predict their performance for fuel purposes and for production of wartime devices such as gas masks. This work was the basis of her PhD thesis ''The physical chemistry of solid organic colloids with special reference to coal'' for which the University of Cambridge awarded her a PhD in 1945. It was also the basis of several papers.


Career and research


Paris

With World War II ending in 1945, Franklin asked Adrienne Weill for help and to let her know of job openings for "a physical chemist who knows very little physical chemistry, but quite a lot about the holes in coal." At a conference in the autumn of 1946, Weill introduced her to Marcel Mathieu, a director of the Centre national de la recherche scientifique (CNRS), the network of institutes that comprises the major part of the scientific research laboratories supported by the French government. This led to her appointment with Jacques Mering at the Laboratoire Central des Services Chimiques de l'État in Paris. She joined the ''labo'' (as referred to by the staff) of Mering on 14 February 1947 as one of the fifteen ''chercheurs'' (researchers). Mering was an X-ray crystallographer who applied X-ray diffraction to the study of rayon and other amorphous substances, in contrast to the thousands of regular crystals that had been studied by this method for many years. He taught her the practical aspects of applying X-ray crystallography to amorphous substances. This presented new challenges in the conduct of experiments and the interpretation of results. Franklin applied them to further problems related to coal and to other carbonaceous materials, in particular the changes to the arrangement of atoms when these are converted to graphite. She published several further papers on this work which has become part of the mainstream of the physics and chemistry of coal and carbon. She coined the terms graphitising and non-graphitising carbon. The coal work was covered in a 1993 monograph, and in the regularly-published textbook ''Chemistry and Physics of Carbon''. Mering continued the study of carbon in various forms, using X-ray diffraction and other methods.


King's College London

In 1950, Franklin was granted a three-year Turner & Newall Fellowship to work at
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
. In January 1951, she started working as a research associate in the Medical Research Council's (MRC) Biophysics Unit, directed by John Randall. She was originally appointed to work on X-ray diffraction of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s and
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
s in solution, but Randall redirected her work to DNA fibres because of new developments in the field, and she was to be the only experienced experimental diffraction researcher at King's at the time. Randall made this reassignment, even before Franklin started working at King's, because of the pioneering work by DNA researcher Maurice Wilkins, and he reassigned Raymond Gosling, the graduate student who had been working with Wilkins, to be her assistant. In 1950, Swiss chemist
Rudolf Signer Rudolf Signer (17 March 1903, Herisau, Switzerland – 1 December 1990, Gümlingen, Switzerland) contributed to the discovery of the DNA double helix. He was a Professor for organic chemistry at the University of Bern from 1935 until 1972 ...
in Berne prepared a highly purified DNA sample from calf
thymus The thymus is a specialized primary lymphoid organ of the immune system. Within the thymus, thymus cell lymphocytes or '' T cells'' mature. T cells are critical to the adaptive immune system, where the body adapts to specific foreign invaders ...
. He freely distributed the DNA sample, later referred to as the Signer DNA, in early May 1950 at the meeting of the Faraday Society in London, and Wilkins was one of the recipients. Even using crude equipment, Wilkins and Gosling had obtained a good-quality diffraction picture of the DNA sample which sparked further interest in this molecule.Professor Raymond Gosling, DNA scientist – obituary
. ''
The Daily Telegraph ''The Daily Telegraph'', known online and elsewhere as ''The Telegraph'', is a national British daily broadsheet newspaper published in London by Telegraph Media Group and distributed across the United Kingdom and internationally. It was f ...
''. 22 May 2015. Retrieved 3 September 2019.
But Randall had not indicated to them that he had asked Franklin to take over both the DNA diffraction work and guidance of Gosling's thesis. It was while Wilkins was away on holiday that Randall, in a letter in December 1950, assured Franklin that "as far as the experimental X-ray effort there would be for the moment only yourself and Gosling." Randall's lack of communication about this reassignment significantly contributed to the well documented friction that developed between Wilkins and Franklin. When Wilkins returned, he handed over the Signer DNA and Gosling to Franklin. Franklin, now working with Gosling, 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, a critical innovation she applied was making the camera chamber that could be controlled for its
humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity dep ...
using different saturated salt solutions. When Wilkins enquired about this improved technique, she replied in terms which offended him as she had "an air of cool superiority". Franklin's habit of intensely looking people in the eye while being concise, impatient and direct unnerved many of her colleagues. In stark contrast, Wilkins was very shy, and slowly calculating in speech while he avoided looking anyone directly in the eye.Elkin p. 45. With the ingenious humidity-controlling camera, Franklin was soon able to produce X-ray images of better quality than those of Wilkins. She immediately discovered that the DNA sample could exist in two forms: at a relative humidity higher than 75%, the DNA fibre became long and thin; when it was drier, it became short and fat. She originally referred to the former as "wet" and the latter as "crystalline." On the structure of the crystalline DNA, Franklin first recorded the analysis in her notebook, which reads: "Evidence for spiral eaning helicalstructure. Straight chain untwisted is highly improbable. Absence of reflections on meridian in χtalline rystallineform suggests spiral structure." An immediate discovery from this was that the phosphate group lies outside of the main DNA chain; she however could not make out whether there could be two or three chains. She presented their data at a lecture in November 1951, in King's College London. In her lecture notes, she wrote the following:
The results suggest a helical structure (which must be very closely packed) containing 2, 3 or 4 co-axial nucleic acid chains per helical unit, and having the phosphate groups near the outside.
Franklin then named " A" and "B" respectively for the "crystalline" and "wet" forms. (The biological functions of A-DNA were discovered only 60 years later.) Because of the intense personality conflict developing between Franklin and Wilkins, Randall divided the work on DNA. Franklin chose the data rich "A" form while Wilkins selected the "B" form.Maddox, p. 155.Wilkins, p. 158. By the end of 1951 it became generally accepted at King's that the B-DNA was a helix, but after she had recorded an asymmetrical image in May 1952, Franklin became unconvinced that the A-DNA was a helix.Wilkins, p. 176. In July 1952, as a practical joke on Wilkins (who frequently expressed his view that both forms of DNA were helical), Franklin and Gosling produced a funeral notice regretting the 'death' of helical A-DNA, which runs:
It is with great regret that we have to announce the death, on Friday 18th July 1952 of DNA helix (crystalline). Death followed a protracted illness which an intensive course of Besselised eferring_to_Bessel_function_that_was_used_to_analyse_the_X-ray_diffraction_patterns.html" ;"title="Bessel_function.html" ;"title="eferring to Bessel function">eferring to Bessel function that was used to analyse the X-ray diffraction patterns">Bessel_function.html" ;"title="eferring to Bessel function">eferring to Bessel function that was used to analyse the X-ray diffraction patternsinjections had failed to relieve. A memorial service will be held next Monday or Tuesday. It is hoped that Dr M H F Wilkins will speak in memory of the late helix. [Signed Rosalind Franklin and Raymond Gosling.]
During 1952, they worked at applying the Patterson function to the X-ray pictures of DNA they had produced. This was a long and labour-intensive approach but would yield significant insight into the structure of the molecule. Franklin was fully committed to experimental data and was sternly against theoretical or model buildings, as she said, "We are not going to speculate, we are going to wait, we are going to let the spots on this photograph tell us what the NAstructure is." The X-ray diffraction pictures, including the landmark '' Photo 51'' taken by Gosling at this time, have been called by John Desmond Bernal as "amongst the most beautiful X-ray photographs of any substance ever taken".Maddox, p. 153. By January 1953, Franklin had reconciled her conflicting data, concluding that both DNA forms had two helices, and had started to write a series of three draft manuscripts, two of which included a double helical DNA backbone (see below). Her two A-DNA manuscripts reached '' Acta Crystallographica'' in
Copenhagen Copenhagen ( or .; da, København ) is the capital and most populous city of Denmark, with a proper population of around 815.000 in the last quarter of 2022; and some 1.370,000 in the urban area; and the wider Copenhagen metropolitan a ...
on 6 March 1953, one day before Crick and Watson had completed their model on B-DNA. She must have mailed them while the Cambridge team was building their model, and certainly had written them before she knew of their work.Maddox, p. 199. 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. The third draft paper was on the B-DNA, dated 17 March 1953, which was discovered years later amongst her papers, by Franklin's Birkbeck colleague,
Aaron Klug Sir Aaron Klug (11 August 1926 – 20 November 2018) was a British biophysicist and chemist. He was a winner of the 1982 Nobel Prize in Chemistry for his development of crystallographic electron microscopy and his structural elucidation of bi ...
. He then published in 1974 an evaluation of the draft's close correlation with the third of the original trio of 25 April 1953 ''Nature'' DNA articles. Klug designed this paper to complement the first article he had written in 1968 defending Franklin's significant contribution to DNA structure. He had written this first article in response to the incomplete picture of Franklin's work depicted in
James Watson James Dewey Watson (born April 6, 1928) is an American molecular biologist, geneticist, and zoologist. In 1953, he co-authored with Francis Crick the academic paper proposing the double helix structure of the DNA molecule. Watson, Crick a ...
's 1968 memoir, '' The Double Helix''. As vividly described by Watson, he travelled to King's on 30 January 1953 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 showed Watson Franklin's DNA X-ray image. Watson, in turn, showed Wilkins a prepublication manuscript by Pauling and Robert Corey, which contained a DNA structure remarkably like their first incorrect model.


Discovery of DNA structure

In February 1953, James Watson and
Francis Crick Francis Harry Compton Crick (8 June 1916 – 28 July 2004) was an English molecular biologist, biophysicist, and neuroscientist. He, James Watson, Rosalind Franklin, and Maurice Wilkins played crucial roles in deciphering the helical stru ...
of the
Cavendish Laboratory The Cavendish Laboratory is the Department of Physics at the University of Cambridge, and is part of the School of Physical Sciences. The laboratory was opened in 1874 on the New Museums Site as a laboratory for experimental physics and is named ...
in
Cambridge University , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
had started to build a molecular model of the B-DNA using data similar to that available to both teams at King's. Based on Franklin's lecture in November 1951 that DNA was helical with either two or three stands, they constructed a triple helix model, which was immediately proven to be flawed. Franklin's research was completed by February 1953, ahead of her move to Birkbeck, and her data was critical. Model building had been applied successfully in the elucidation of the structure of the
alpha helix The alpha helix (α-helix) is a common motif in the secondary structure of proteins and is a right hand- helix conformation in which every backbone N−H group hydrogen bonds to the backbone C=O group of the amino acid located four residues ...
by Linus Pauling in 1951, but Franklin was opposed to prematurely building theoretical models, until sufficient data were obtained to properly guide the model building. She took the view that building a model was to be undertaken only after enough of the structure was known. Her conviction was justified by the fact that Pauling and Corey also came up in the late 1952 (published in February 1953) with an erroneous triple helix model. Ever cautious, she wanted to eliminate misleading possibilities. Photographs of her Birkbeck work table show that she routinely used small molecular models, although certainly not ones on the grand scale successfully used at Cambridge for DNA. In the middle of February 1953, Crick's thesis advisor,
Max Perutz Max Ferdinand Perutz (19 May 1914 – 6 February 2002) was an Austrian-born British molecular biologist, who shared the 1962 Nobel Prize for Chemistry with John Kendrew, for their studies of the structures of haemoglobin and myoglobin. He went ...
, 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". However, they knew they must complete their model before they could be certain. 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". This was also one day after Franklin's two A-DNA papers had reached ''Acta Crystallographica''. Wilkins came to see the model the following week, according to Franklin's biographer
Brenda Maddox Brenda, Lady Maddox ( Murphy; February 24, 1932 – June 16, 2019) was an American writer and biographer, who spent most of her adult life living and working in the UK, from 1959 until her death. She is best known for her biographies, includin ...
, on 12 March, and allegedly informed Gosling on his return to King's.Maddox, p. 207. One of the most critical and overlooked moments in DNA research was how and when Franklin realised and conceded that B-DNA was a double helical molecule. When Klug first examined Franklin's documents after her death, he initially came to an impression that Franklin was not convinced of the double helical nature until the knowledge of the Cambridge model. But he later discovered the original draft of the manuscript (dated 17 March 1953) from which it became clear that Franklin had already resolved the correct structure. The news of Watson–Crick model reached King's the next day, 18 March, suggesting that Franklin would have learned of it much later since she had moved to Birkbeck. Further scrutiny of her notebook revealed that Franklin had already thought of the helical structure for B-DNA in February 1953 but was not sure of the number of strands, as she wrote: "Evidence for 2-chain (or 1-chain helix)."Olby, p. 418. Her conclusion on the helical nature was evident, though she failed to understand the complete organisation of the DNA strands as the possibility two strands running in opposite directions did not occur to her. Towards the end of February she began to work out the indications of double strands, as she noted: "Structure B does not fit single helical theory, even for low layer-lines." It soon dawned to her that the B-DNA and A-DNA were structurally similar, and perceived A-DNA as an "unwound version" of B-DNA. She and Gosling wrote a five-paged manuscript on 17 March titled "A Note on Molecular Configuration of Sodium Thymonucleate." After the Watson–Crick model was known, there appeared to be only one (hand-written) modification after the typeset at the end of the text which states that their data was consistent with the model, and appeared as such in the trio of 25 April 1953 ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
'' articles; the other modification being a deletion of "A Note on" from the title. eproduction with interpretation: /ref> Weeks later, on 10 April, Franklin wrote to Crick for permission to see their model. Franklin retained her scepticism for premature model building even after seeing the Watson–Crick model, and remained unimpressed. She is reported to have commented, "It's very pretty, but how are they going to prove it?" As an experimental scientist, Franklin seems to have been interested in producing far greater evidence before publishing-as-proven a proposed model. Accordingly, her response to the Watson–Crick model was in keeping with her cautious approach to science. Crick and Watson then published their model in ''Nature'' on 25 April 1953, in an article describing the double-helical structure of DNA with only a footnote acknowledging "having been stimulated by a general knowledge of Franklin and Wilkins' ‘unpublished’ contribution.” Actually, although it was the bare minimum, they had just enough specific knowledge of Franklin and Gosling's data upon which to base their model. As a result of a deal struck by the two laboratory directors, articles by Wilkins and Franklin, which included their X-ray diffraction data, were modified and then published second and third in the same issue of ''Nature'', seemingly only in support of the Crick and Watson theoretical paper which proposed a model for the B-DNA. 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.


Birkbeck College

Franklin left King's College London in mid-March 1953 for
Birkbeck College , mottoeng = Advice comes over nightTranslation used by Birkbeck. , established = , type = Public research university , endowment = £4.3 m (2014) , budget = £10 ...
, in a move that had been planned for some time and that she described (in a letter to Adrienne Weill in Paris) as "moving from a palace to the slums ... but pleasanter all the same". She was recruited by physics department chair John Desmond Bernal, a crystallographer who was a communist, known for promoting female crystallographers. Her new laboratories were housed in 21 Torrington Square, one of a pair of dilapidated and cramped Georgian houses containing several different departments; Franklin frequently took Bernal to task over the careless attitudes of some of the other laboratory staff, notably after workers in the pharmacy department flooded her first-floor laboratory with water on one occasion. Despite the parting words of Bernal to stop her interest in nucleic acids, she helped Gosling to finish his thesis, although she was no longer his official supervisor. Together they published the first evidence of double helix in the A form of DNA in the 25 July issue of ''Nature''. At the end of 1954, Bernal secured funding for Franklin from the
Agricultural Research Council The Agricultural and Food Research Council (AFRC) was a British Research Council responsible for funding and managing scientific and technological developments in farming and horticulture. History The AFRC was formed in 1983 from its predecessor, ...
(ARC), which enabled her to work as a senior scientist supervising her own research group.Brown, pp. 356–357. John Finch, a physics student from
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
, subsequently joined Franklin's group, followed by
Kenneth Holmes Kenneth Charles Holmes FRS (1934 – 2 November 2021) was a British molecular biologist. He was born in Hammersmith, London. He was a former colleague of Rosalind Franklin at Birkbeck College with Aaron Klug and John Finch and moved to the Labo ...
, a Cambridge graduate, in July 1955. Despite the ARC funding, Franklin wrote to Bernal that the existing facilities remained highly unsuited for conducting research "...my desk and lab are on the fourth floor, my X-ray tube in the basement, and I am responsible for the work of four people distributed over the basement, first and second floors on two different staircases."


RNA research

Franklin continued to explore another major nucleic acid, RNA, a molecule equally central to life as DNA. She again used X-ray crystallography to study the structure of the tobacco mosaic virus (TMV), an
RNA virus An RNA virus is a virusother than a retrovirusthat has ribonucleic acid ( RNA) as its genetic material. The nucleic acid is usually single-stranded RNA (ssRNA) but it may be double-stranded (dsRNA). Notable human diseases caused by RNA virus ...
. Her meeting with Aaron Klug in early 1954 led to a longstanding and successful collaboration. Klug had just then earned his PhD from
Trinity College, Cambridge Trinity College is a constituent college of the University of Cambridge. Founded in 1546 by King Henry VIII, Trinity is one of the largest Cambridge colleges, with the largest financial endowment of any college at either Cambridge or Oxford. ...
, and joined Birkbeck in late 1953. In 1955 Franklin published her first major works on TMV in ''Nature'', in which she described that all TMV virus particles were of the same length. This was in direct contradiction to the ideas of the eminent virologist
Norman Pirie Norman Wingate (Bill) Pirie FRS (1 July 1907 – 29 March 1997), was a British biochemist and virologist who, along with Frederick Bawden, discovered that a virus can be crystallized by isolating tomato bushy stunt virus in 1936. This was an ...
, though her observation ultimately proved correct. Franklin assigned the study of the complete structure of TMV to her PhD student Holmes. They soon discovered (published in 1956) that the covering of TMV was protein molecules arranged in helices. Her colleague Klug worked on spherical viruses with his student Finch, with Franklin coordinating and overseeing the work. As a team, from 1956 they started publishing seminal works on TMV, cucumber virus 4 and
turnip yellow mosaic virus ''Turnip yellow mosaic virus'' (TYMV) is an isometric ''Tymovirus'' of the family ''Tymoviridae''. Its host range is confined almost entirely to plants in the genus '' Brassica'' in western Europe, which includes cabbages, cauliflower and br ...
. 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. The Birkbeck team members continued working on RNA viruses affecting several plants, including potato, turnip, tomato and pea. In 1955 the team was joined by an American post-doctoral student Donald Caspar. He worked on the precise location of RNA molecules in TMV. In 1956 he and Franklin published individual but complementary papers in the 10 March issue of ''Nature'', in which they showed that the RNA in TMV is wound along the inner surface of the hollow virus. Caspar was not an enthusiastic writer, and Franklin had to write the entire manuscript for him. Her research grant from ARC expired at the end of 1957, and she was never given the full salary proposed by Birkbeck. After Bernal requested ARC chairman
Lord Rothschild Baron Rothschild, of Tring in the County of Hertfordshire, is a title in the Peerage of the United Kingdom. It was created in 1885 for Sir Nathan Rothschild, 2nd Baronet, a member of the Rothschild banking family. He was the first Jewish m ...
, she was given a one-year extension ending in March 1958. Expo 58, the first major international fair after World War II, was to be held in Brussels in 1958. Franklin was invited to make a five-foot high model of TMV, which she started in 1957. Her materials included table tennis balls and plastic bicycle handlebar grips. The Brussels world's fair, with an exhibit of her virus model at the International Science Pavilion, opened on 17 April, one day after she died.


Polio virus

In 1956, Franklin visited the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant un ...
, where colleagues suggested her group research the polio virus. In 1957 she applied for a grant from the United States Public Health Service of the
National Institutes of Health The National Institutes of Health, commonly referred to as NIH (with each letter pronounced individually), is the primary agency of the United States government responsible for biomedical and public health research. It was founded in the lat ...
, which approved £10,000 (equivalent to £ in ) for three years, the largest fund ever received at Birkbeck. In her grant application, Franklin mentioned her new interest in animal virus research. She obtained Bernal's consent in July 1957, though serious concerns were raised after she disclosed her intentions to research live, instead of killed, polio virus at Birkbeck. Eventually, Bernal arranged for the virus to be safely stored at the London School of Hygiene and Tropical Medicine during the group's research. With her group, Franklin then commenced deciphering the structure of the polio virus while it was in a crystalline state. She attempted to mount the virus crystals in capillary tubes for X-ray studies, but was forced to end her work due to her rapidly failing health. After Franklin's death, Klug succeeded her as group leader, and he, Finch and Holmes continued researching the structure of the polio virus. They eventually succeeded in obtaining extremely detailed X-ray images of the virus. In June 1959, Klug and Finch published the group's findings, revealing the polio virus to have icosahedral symmetry, and in the same paper suggested the possibility for all spherical viruses to possess the same symmetry, as it permitted the greatest possible number (60) of identical structural units. The team moved to the Laboratory of Molecular Biology, Cambridge, in 1962, and the old Torrington Square laboratories were demolished four years later, in May 1966.


Personal life

Franklin was best described as an agnostic. Her lack of religious faith apparently did not stem from anyone's influence, rather from her own line of thinking. She developed her scepticism as a young child. Her mother recalled that she refused to believe in the existence of God, and remarked, "Well, anyhow, how do you know He isn't She?" She later made her position clear, now based on her scientific experience, and wrote to her father in 1940: However, she did not abandon Jewish traditions. As the only Jewish student at Lindores School, she had Hebrew lessons on her own while her friends went to church. She joined the Jewish Society while in her first term at Cambridge, out of respect of her grandfather's request. Franklin confided to her sister that she was "always consciously a Jew". Franklin loved travelling abroad, particularly trekking. She first "qualified" at Christmas 1929 for a vacation at Menton, France, where her grandfather went to escape the English winter. Her family frequently spent vacations in Wales or Cornwall. A trip to France in 1938 gave her a lasting love for France and its language. She considered the French lifestyle at that time as "vastly superior to that of English".Polcovar, p. 33. In contrast, she described English people as having "vacant stupid faces and childlike complacency". Her family was almost stuck in Norway in 1939, as World War II was declared on their way home. In another instance, she trekked the French Alps with Jean Kerslake in 1946, which almost cost her her life. She slipped off a slope, and was barely rescued. But she wrote to her mother, "I am quite sure I could wander happily in France forever. I love the people, the country and the food."Polcovar, p. 41. She made several professional trips to the United States, and was particularly jovial among her American friends and constantly displayed her sense of humour. William Ginoza of the University of California, Los Angeles, later recalled that she was the opposite of Watson's description of her, and as Maddox comments, Americans enjoyed her "sunny side". In his book ''The Double Helix'', Watson provides his first-person account of the search for and discovery of DNA. He paints a sympathetic but sometimes critical portrait of Franklin. He praises her intellect and scientific acumen, but portrays her as difficult to work with and careless with her appearance. After introducing her in the book as "Rosalind", he writes that he and his male colleagues usually referred to her as "Rosy", the name people at King's College London used behind her back. She did not want to be called by that name because she had a great-aunt Rosy. In the family, she was called "Ros". To others, she was simply "Rosalind". She made it clear to an American visiting friend, Dorothea Raacke, while sitting with her at Crick's table in The Eagle, Cambridge, The Eagle pub in Cambridge: Raacke asked her how she was to be called and she replied "I'm afraid it will have to be Rosalind", adding "Most definitely not ''Rosy''."Maddox, p. 288. She often expressed her political views. She initially blamed Winston Churchill for inciting the war, but later admired him for his speeches. She actively supported Professor John Ryle (physician), John Ryle as an independent candidate for parliament in the 1940 Cambridge University by-election, but he was unsuccessful.Glynn, p. 52. She did not seem to have an intimate relationship with anyone, and always kept her deepest personal feelings to herself. After her younger days, she avoided close friendship with the opposite sex. In her later years, Evi Ellis, who had shared her bedroom when a child refugee and who was then married to Ernst Wohlgemuth and had moved to Notting Hill from Chicago, tried matchmaking her with Ralph Miliband but failed. Franklin once told Evi that her flatmate asked her for a drink, but she did not understand the intention. She was quite infatuated by her French mentor Mering, who had a wife and a mistress. Mering also admitted that he was captivated by her "intelligence and beauty". According to Anne Sayre, Franklin did confess her feeling for Mering when she was undergoing a second surgery, but Maddox reported that the family denied this. Mering wept when he visited her later, and destroyed all her letters after her death. Her closest personal affair was probably with her once post-doctoral student Donald Caspar. In 1956, she visited him at his home in Colorado after her tour to
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant un ...
, and she was known to remark later that Caspar was one "she might have loved, might have married". In her letter to Sayre, she described him as "an ideal match".


Illness, death, and burial

In mid-1956, while on a work-related trip to the United States, Franklin first began to suspect a health problem. While in New York she found difficulty in zipping her skirt; her stomach had bulged. Back in London she consulted Mair Livingstone, who asked her, "You're not pregnant?" to which she retorted, "I wish I were." Her case was marked "URGENT". An operation on 4 September of the same year revealed two tumours in her abdomen. After this period and other periods of hospitalisation, 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 six more in 1957. At the end of 1957, Franklin again fell ill and she was admitted to the Royal Marsden Hospital. On 2 December, she made her will. She named her three brothers as executors and made her colleague Aaron Klug the principal beneficiary, who would receive £3,000 and her Austin car. Of her other friends, Mair Livingstone would get £2,000, Anne Piper £1,000, and her nurse Miss Griffith £250. The remainder of the estate was to be used for charities. She returned to work in January 1958, and was also given a promotion to Research Associate in Biophysics on 25 February. She fell ill again on 30 March, and she died on 16 April 1958, in Chelsea, London, Chelsea, London, of bronchopneumonia, secondary carcinomatosis, and
ovarian cancer Ovarian cancer is a cancerous tumor of an ovary. It may originate from the ovary itself or more commonly from communicating nearby structures such as fallopian tubes or the inner lining of the abdomen. The ovary is made up of three different ...
. Health effects of radiation, Exposure to X-ray radiation is sometimes considered to be a possible factor in her illness.Along with genetic predisposition; opinion of CSU's Lynne Osman Elkin; see also March 2003 ''Physics Today'' Other members of her family have died of cancer, and the incidence of gynaecological cancer is known to be disproportionately high among Ashkenazi Jews. Her death certificate states: ''A Research Scientist, Spinster, Daughter of Ellis Arthur Franklin, a Banker.'' She was interred on 17 April 1958 in the family plot at Willesden United Synagogue Cemetery at Beaconsfield Road in London Borough of Brent. The inscription on her tombstone reads:
IN MEMORY OF
ROSALIND ELSIE FRANKLIN
מ' רחל בת ר' יהודה [Rochel/Rachel daughter of Yehuda, her father's Hebrew name]
DEARLY LOVED ELDER DAUGHTER OF
ELLIS AND MURIEL FRANKLIN
25TH JULY 1920 – 16TH APRIL 1958
SCIENTIST
HER RESEARCH AND DISCOVERIES ON
VIRUSES REMAIN OF LASTING BENEFIT
TO MANKIND
ת נ צ ב ה [Hebrew initials for "her soul shall be bound in the bundle of life"]
Franklin's will was proven on 30 June, with her estate assessed for probate at £11,278 10s. 9d. (equivalent to £ in ).


Controversies after death


Alleged sexism toward Franklin

Anne Sayre, Franklin's friend and one of her biographers, says in her 1975 book, ''Rosalind Franklin and DNA: '' "In 1951 ... King's College London as an institution, was not distinguished for the welcome that it offered to women ... Rosalind ... was unused to ''purdah'' [a religious and social institution of female seclusion] ... there was one other female on the laboratory staff". The molecular biology, molecular biologist Andrzej Stasiak notes: "Sayre's book became widely cited in feminist circles for exposing rampant sexism in science." Farooq Hussain says: "there were seven women in the biophysics department ... Jean Hanson became an FRS, Dame Honor B. Fell, Director of Strangeways Laboratory, supervised the biologists". Maddox states: "John Randall (physicist), Randall ... did have many women on his staff ... they found him ... sympathetic and helpful."Maddox, p. 135. Sayre asserts 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". However, Elkin claims that most of the Medical Research Council (UK), MRC group (including Franklin) typically ate lunch together in the mixed dining room discussed below. And Maddox says, of Randall: "He liked to see his flock, men and women, come together for morning coffee, and at lunch in the joint dining room, where he ate with them nearly every day." Francis Crick also commented that "her colleagues treated men and women scientists alike". Sayre also discusses at length Franklin's struggle in pursuing science, particularly her father's concern about women in academic professions. This account had led to accusations of sexism in regard to Ellis Franklin's attitude to his daughter. A good deal of information explicitly claims that he strongly opposed her entering Newnham College. The Public Broadcasting Service (PBS) biography of Franklin goes further, stating that he refused to pay her fees, and that an aunt stepped in to do that for her. Her sister, Jenifer Glynn, has stated that those stories are myths, and that her parents fully supported Franklin's entire career. Sexism is said to pervade the memoir of one peer, James Watson, in his book ''The Double Helix'', published 10 years after Franklin's death and after Watson had returned from Cambridge to Harvard. His Cambridge colleague, Peter Pauling, wrote in a letter, "Morris [''sic''] Wilkins is supposed to be doing this work; Miss Franklin is evidently a fool." Crick acknowledges later, "I'm afraid we always used to adopt – let's say, a ''patronizing'' attitude towards her." Glynn accuses Sayre of erroneously making her sister a feminist heroine, and sees Watson's ''The Double Helix'' as the root of what she calls the "Rosalind Industry". She conjectures that the stories of alleged sexism would "have embarrassed her [Rosalind Franklin] almost as much as Watson's account would have upset her", and declared that "she [Rosalind] was never a feminist." Klug and Crick have also concurred that Franklin was definitely not a feminist. Franklin's letter to her parents in January 1939 is often taken as reflecting her own prejudiced attitude, and the claim that she was "not immune to the sexism rampant in these circles". In the letter, she remarked that one lecturer was "very good, though female". Maddox maintains that was a circumstantial comment rather than an example of gender bias, and that it was a expression of admiration because, at the time, woman teachers of science were a rarity. In fact, Maddox says, Franklin laughed at men who were embarrassed by the appointment of the first female professor, Dorothy Garrod.


Contribution to the model/structure of DNA

Rosalind Franklin's first important contributions to the model popularised by Crick and Watson was her lecture at the seminar in November 1951, where she presented to those present, among them Watson, the two forms of the molecule, type A and type B, her position being that the phosphate units are located in the external part of the molecule. She also specified the amount of water to be found in the molecule in accordance with other parts of it, data that have considerable importance for the stability of the molecule. Franklin was the first to discover and formulate these facts, which in fact constituted the basis for all later attempts to build a model of the molecule. However, Watson, at the time ignorant of the chemistry, failed to comprehend the crucial information, and this led to the construction of a wrong three-helical model. The other contribution included an X-ray photograph of B-DNA (called '' Photo 51'') taken by Franklin's student Gosling that was briefly shown to Watson by Wilkins in January 1953,Maddox, p. 196. and a report written for an MRC biophysics committee visit to King's in December 1952 which was shown by Perutz at the Cavendish Laboratory to both Crick and Watson. This MRC report contained data from the King's group, including some of Franklin's and Gosling's work, and was given to Crick – who was working on his thesis on haemoglobin structure – by his thesis supervisor Perutz, a member of the visiting committee. Sayre's biography of Franklin contains a story alleging that the photograph 51 in question was shown to Watson by Wilkins without Franklin's permission, and that this constituted a case of bad science ethics. Others dispute this story, asserting that Wilkins had been given photograph 51 by Franklin's PhD student Gosling because she was leaving King's to work at Birkbeck, and there was allegedly nothing untoward in this transfer of data to Wilkins because Director Randall had insisted that all DNA work belonged exclusively to King's and had instructed Franklin in a letter to even stop working on it and submit her data. Also, it was implied by Horace Freeland Judson, that Maurice Wilkins had taken the photograph out of Franklin's drawer, but this is also said to be incorrect. Likewise, Perutz saw "no harm" in showing an MRC report containing the conclusions of Franklin and Gosling's X-ray data analysis to Crick, since it had not been marked as confidential, although "The report was not expected to reach outside eyes". 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 and to post a general statement in ''Science (journal), Science'' excusing himself on the basis of being "inexperienced and casual in administrative matters". Perutz also claimed that the MRC information was already made available to the Cambridge team when Watson had attended Franklin's seminar in November 1951. A preliminary version of much of the important material contained in the 1952 December MRC report had been presented by Franklin in a talk she had given in November 1951, which Watson had attended but not understood. The Perutz letter was as said one of three letters, published with letters by Wilkins and Watson, which discussed their various contributions. Watson clarified the importance of the data obtained from the MRC report as he had not recorded these data while attending Franklin's lecture in 1951. The upshot of all this was that when Crick and Watson started to build their model in February 1953 they were working with critical parameters that had been determined by Franklin in 1951, and which she and Gosling had significantly refined in 1952, as well as with published data and other very similar data to those available at King's. It was generally believed that Franklin was never aware that her work had been used during construction of the model, but Gosling asserted in his 2013 interview that, "Yes. Oh, she did know about that."


Recognition of her contribution to the model of DNA

Upon the completion of their model, Crick and Watson had invited Wilkins to be a co-author of their paper describing the structure. Wilkins turned down this offer, as he had taken no part in building the model. He later expressed regret that greater discussion of co-authorship had not taken place as this might have helped to clarify the contribution the work at King's had made to the discovery. There is no doubt that Franklin's experimental data were used by Crick and Watson to build their model of DNA in 1953. Some, including Maddox, have explained this citation omission by suggesting that it may be a question of circumstance, because it would have been very difficult to cite the unpublished work from the MRC report they had seen. Indeed, a clear timely acknowledgment would have been awkward, given the unorthodox manner in which data were transferred from King's to Cambridge. However, methods were available. Watson and Crick could have cited the MRC report as a personal communication or else cited the ''Acta ''articles in press, or most easily, the third ''Nature'' paper that they knew was in press. One of the most important accomplishments of Maddox's widely acclaimed biography is that Maddox made a well-received case for inadequate acknowledgement. "Such acknowledgement as they gave her was very muted and always coupled with the name of Wilkins". Fifteen years after the fact, the first clear recitation of Franklin's contribution appeared as it permeated Watson's account, ''The Double Helix'', although it was buried under descriptions of Watson's (often quite negative) regard towards Franklin during the period of their work on DNA. This attitude is epitomized in the confrontation between Watson and Franklin over a preprint of Pauling's mistaken DNA manuscript. Watson's words impelled Sayre to write her rebuttal, in which the entire chapter nine, "Winner Take All" has the structure of a legal brief dissecting and analyzing the topic of acknowledgement. Sayre's early analysis was often ignored because of perceived feminist overtones in her book. Watson and Crick did not cite the X-ray diffraction work of Wilkins and Franklin in their original paper, though they admit having "been stimulated by a knowledge of the general nature of the unpublished experimental results and ideas of Dr. M. H. F. Wilkins, Dr. R. E. Franklin and their co-workers at King's College London". In fact, Watson and Crick cited no experimental data at all in support of their model. Franklin and Gosling's publication of the DNA X-ray image, in the same issue of ''Nature'', served as the principal evidence:
Thus our general ideas are not inconsistent with the model proposed by Watson and Crick in the preceding communication.


Nobel Prize

Franklin was never nominated for a Nobel Prize. Her work was a crucial part in the discovery of DNA's structure, which along with subsequent related work led to Francis Crick, James Watson, and Maurice Wilkins being awarded a Nobel Prize in 1962. She had died in 1958, and during her lifetime the DNA structure was not considered to be fully proven. It took Wilkins and his colleagues about seven years to collect enough data to prove and refine the proposed DNA structure. Moreover, its biological significance, as proposed by Watson and Crick, was not established. General acceptance for the DNA double helix and its function did not start until late in the 1950s, leading to Nobel nominations in 1960, 1961, and 1962 for Nobel Prize in Physiology or Medicine, and in 1962 for Nobel Prize in Chemistry. The first breakthrough was from Matthew Meselson and Franklin Stahl in 1958, who experimentally showed the DNA replication of a bacterium ''Escherichia coli''. In what is now known as the Meselson–Stahl experiment, DNA was found to replicate into two double-stranded helices, with each helix having one of the original DNA strands. This DNA replication was firmly established by 1961 after further demonstration in other species, and of the stepwise chemical reaction. According to the 1961 Crick–Monod letter, this experimental proof, along with Wilkins having initiated the DNA diffraction work, were the reasons why Crick felt that Wilkins should be included in the DNA Nobel Prize. In 1962 the Nobel Prize was subsequently awarded to Crick, Watson, and Wilkins. Nobel rules now prohibit posthumous nominations (though this statute was not formally in effect until 1974) or splitting of Prizes more than three ways. The award was for their body of work on nucleic acids and not exclusively for the discovery of the structure of DNA. By the time of the award Wilkins had been working on the structure of DNA for more than 10 years, and had done much to confirm the Watson–Crick model. Crick had been working on the genetic code at Cambridge and Watson had worked on RNA for some years.Wilkins, p. 243. Watson has suggested that ideally Wilkins and Franklin would have been awarded the Nobel Prize in Chemistry. Pauling, who received the Nobel Peace Prize that year, believed and earlier warned the Nobel Committee in 1960 that "it might well be premature to make an award of a Prize to Watson and Crick, because of existing uncertainty about the detailed structure of nucleic acid. I myself feel that it is likely that the general nature of the Watson-Crick structure is correct, but that there is doubt about details." He was partly right as an alternative of Watson-Crick base pairing, called the Hoogsteen base pairing that can form triple DNA strand, was discovered by Karst Hoogsteen in 1963. Aaron Klug, Franklin's colleague and principal beneficiary in her will, was the sole winner of the Nobel Prize in Chemistry 1982, "for his development of crystallographic electron microscopy and his structural elucidation of biologically important nucleic acid-protein complexes". This work was exactly what Franklin had started and which she introduced to Klug, and it is highly plausible that, were she alive, she would have shared the Nobel Prize.


Awards and honours


Posthumous recognition

* 1982, Iota Sigma Pi designated Franklin a National Honorary Member. * 1984, St Paul's Girls School established the Rosalind Franklin Technology Centre. * 1992, English Heritage placed a blue plaque commemorating Franklin on the building in Drayton Gardens, London, where she lived until her death. * 1993,
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
renamed the Orchard Residence at its Hampstead Campus as Rosalind Franklin Hall. * 1993,
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
placed a blue plaque on its outside wall bearing the inscription: "R. E. Franklin, R. G. Gosling, A. R. Stokes, M. H. F. Wilkins, H. R. Wilson – King's College London – DNA – X-ray diffraction studies – 1953." * 1995,
Newnham College, Cambridge Newnham College is a women's constituent college of the University of Cambridge. The college was founded in 1871 by a group organising Lectures for Ladies, members of which included philosopher Henry Sidgwick and suffragist campaigner Millic ...
opened a graduate residence named Rosalind Franklin Building, and put a bust of her in its garden. * 1997, Birkbeck, University of London School of Crystallography opened the Rosalind Franklin Laboratory. * 1997, a newly discovered asteroid was named 9241 Rosfranklin. * 1998, National Portrait Gallery (London), National Portrait Gallery in London added Rosalind Franklin's portrait next to those of Francis Crick, James Watson and Maurice Wilkins. * 1999, the Institute of Physics at Portland Place, London, renamed its theatre as Franklin Lecture Theatre. * 2000,
King's College London King's College London (informally King's or KCL) is a public research university located in London, England. King's was established by royal charter in 1829 under the patronage of King George IV and the Duke of Wellington. In 1836, King's ...
opened the Franklin–Wilkins Building in honour of Franklin's and Wilkins's work at the college. * 2000, We the Curious (formally @Bristol) features the Rosalind Franklin Room. * 2001, the American National Cancer Institute established the Rosalind E. Franklin Award for women in cancer research. * 2002, the University of Groningen, supported by the European Union, launched the Rosalind Franklin Fellowship to encourage women researchers to become full university professors. * 2003, the Royal Society established the Rosalind Franklin Award (officially the Royal Society Rosalind Franklin Award and Lecture) for an outstanding contribution to any area of natural science, engineering or technology. The award consists of a silver-coated medal and a grant of £30,000. * 2003, the Royal Society of Chemistry declared King's College London as "National Historic Chemical Landmark" and placed a plaque on the wall near the entrance of the building, with the inscription: "Near this site Rosalind Franklin, Maurice Wilkins, Raymond Gosling, Alexander Stokes and Herbert Wilson performed experiments that led to the discovery of the structure of DNA. This work revolutionised our understanding of the chemistry behind life itself." * 2004, Finch University of Health Sciences/The Chicago Medical School, located in North Chicago, Illinois, North Chicago, Illinois, USA changed its name to the Rosalind Franklin University of Medicine and Science. It also adopted a new motto "Life in Discovery", and ''Photo 51'' as its logo. * 2004, the Gruber Foundation started the Rosalind Franklin Young Investigator Award for two female geneticists from all over the world. It carries an annual fund of $25,000, each award is for three years, and selection is made by a joint committee appointed by the Genetics Society of America and the American Society of Human Genetics. * 2004, the Advanced Photon Source (APS) and the APS Users Organization (APSUO) started the APSUO Rosalind Franklin Young Investigator Award for young scientists who made contributions through the APS. * 2005, the DNA sculpture (donated by James Watson) outside Clare College, Cambridge's Memorial Court incorporates the words ''"The double helix model was supported by the work of Rosalind Franklin and Maurice Wilkins."'' * 2005, the Ovarian Cancer Research Alliance, based in Florida, US, established an annual award the Rosalind Franklin Prize for Excellence in Ovarian Cancer Research. * 2006, the Rosalind Franklin Society was established in New York by Mary Ann Liebert. The Society aims to recognise, foster, and advance the important contributions of women in the life sciences and affiliated disciplines. * 2008, Columbia University awarded an honorary Louisa Gross Horwitz Prize to Franklin, "for her seminal contributions to the discovery of the structure of DNA". * 2008, the Institute of Physics established a biennial award the Rosalind Franklin Medal and Prize. * 2012, the bioinformatics education software platform ''Rosalind (education), Rosalind'' was named in honour of Franklin. * 2012, The Rosalind Franklin Building was opened at Nottingham Trent University. * 2013, Google honoured Rosalind Franklin with a Google Doodle, doodle, showing her gazing at a double helix structure of DNA with an X-ray of ''Photo 51'' beyond it. * 2013, a plaque was placed on the wall of The Eagle (pub), The Eagle pub in Cambridge commemorating Franklin's contribution to the discovery of the structure of DNA, on the sixtieth anniversary of Crick and Watson's announcement in the pub. * 2014, the Rosalind Franklin Award for Leadership in Industrial Biotechnology was established by Biotechnology Industry Organization (Biotechnology Innovation Organization since 2016) in collaboration with the Rosalind Franklin Society, for an outstanding woman in the field of industrial biotechnology and bioprocessing. * 2014, the Rosalind Franklin University of Medicine and Science unveiled a bronze statue of Franklin, created by Julie Rotblatt-Amrany, near its front entrance. * 2014, the Rosalind Franklin STEM Elementary was opened in Pasco, Washington, Pasco, Washington, the first science, technology, engineering, and math (STEM) elementary school in the district. * 2014, the University of Wolverhampton opened its new laboratory building named the Rosalind Franklin Science Building. * 2015, Newnham College Boat Club, Cambridge, launched a new racing VIII, naming it the ''Rosalind Franklin''. * 2015, the Rosalind Franklin Appathon was launched by University College London as a national app competition for women in STEMM (science, technology, engineering, maths and medicine). * 2015, a high performance computing and cloud facility in London was named ''Rosalind''. * 2016, the British Humanist Association added the Rosalind Franklin Lecture to its annual lecture series, aimed to explore and celebrate the contribution of women towards the promotion and advancement of humanism. * 2016, the Rosalind Franklin Prize and Tech Day was held on 23 February in London, organised by University College London, i-sense, UCL Enterprise, the London Centre for Nanotechnology and the UCL Athena SWAN Charter. * 2017, DSM (company), DSM opened the Rosalind Franklin Biotechnology Center in Delft, the Netherlands. * 2017, Historic England listed building, gave a heritage listing, at Grade II, to Franklin's tomb at Willesden Jewish Cemetery on the grounds of it being of "special architectural or historic interest". Historic England said that "the tomb commemorates the life and achievements of Rosalind Franklin, a scientist of exceptional distinction, whose pioneering work helped lay the foundations of molecular biology; Franklin’s X-ray observation of DNA contributed to the discovery of its helical structure." * 2018, the Rosalind Franklin Institute, an autonomous medical research centre under the join venture of 10 universities and funded by the United Kingdom Research and Innovation, was launched at the Harwell Science and Innovation Campus on 6 June, and was officially opened on 29 September 2021. * 2019, the European Space Agency (ESA) named its ExoMars rover ''Rosalind Franklin (rover), Rosalind Franklin''. * 2019, the University of Portsmouth announced that it changed the name James Watson Halls to Rosalind Franklin Halls from 2 September. * 2020, Franklin was selected for the Time Person of the Year, ''Time'' 100 Women of the Year, for 1953. * 2020, the UK Royal Mint released a 50-pence coin in honour of the hundredth anniversary of Franklin's birth on 25 July. It features a stylized version of ''Photo 51''. * 2020, South Norfolk Council renamed a road on the Norwich Research Park in her honour in July 2020. The road is home to the Quadram Institute and the University of East Anglia's Bob Champion Research and Education Building. * 2020, Trinity College Dublin announced that its library had previously held forty busts, all of whom were of men, was commissioning four new busts of women one of whom would be Franklin. * 2020, Aston Medical School instituted an annual competition for medical students named the Rosalind Franklin Essay Prize, funded by its alumni and Rosalind's nephew, Daniel Franklin, executive and diplomatic editor of ''The Economist.'' * 2021, a bronze Tondo (art), tondo of Rosalind Franklin was placed on Hampstead Manor and unveiled on 15 March. * On 30 June 2021, a satellite named after her (ÑuSat 19 or "Rosalind", COSPAR 2021-059AC) was launched into space. * 2021, the Rosalind Franklin laboratory was opened in the Royal Leamington Spa, Warwickshire, on 13 July as the largest laboratory for COVID-19 testing under the UK Health Security Agency and National Health Service (England), National Health Service Test and Trace network, and supported by the University of Warwick. * 2022, the new bacterial genus, ''Franklinella'', in the family Comamonadaceae, was described in her honour.


Cultural references

Franklin's part in the discovery of the nature of DNA was shown in the 1987 TV Movie ''Life Story (TV film), Life Story'', starring Juliet Stevenson as Franklin, and with Tim Pigott-Smith as Crick, Alan Howard (actor), Alan Howard as Wilkins and Jeff Goldblum as Watson. This movie portrayed Franklin as somewhat stern, but also alleged that Watson and Crick did use a lot of her work to do theirs. A 56-minute documentary of the life and scientific contributions of Franklin, ''DNA – Secret of Photo 51'', was broadcast in 2003 on PBS Nova, PBS ''Nova''. Narrated by Barbara Flynn, the program features interviews with Wilkins, Gosling, Klug, Maddox, including Franklin's friends Vittorio Luzzati, Caspar, Anne Piper, and Sue Richley. The UK version produced by BBC is titled ''Rosalind Franklin: DNA's Dark Lady''. The first episode of another PBS documentary serial, ''DNA'', was aired on 4 January 2004. The episode titled ''The Secret of Life'' centres much around the contributions of Franklin. Narrated by Jeff Goldblum, it features Watson, Wilkins, Gosling and Peter Pauling (son of Linus Pauling). A play entitled ''Rosalind: A Question of Life'' was written by Deborah Gearing to mark the work of Franklin, and was first performed on 1 November 2005 at the Birmingham Repertory Theatre, and published by Oberon Books in 2006. Another play, ''Photograph 51 (play), Photograph 51'' by Anna Ziegler (playwright), Anna Ziegler, published in 2011, has been produced at several places in the US and in late 2015 was put on at the Noel Coward Theatre, London, with Nicole Kidman playing Franklin. Ziegler's version of the 1951–53 'race' for the structure of DNA sometimes emphasizes the pivotal role of Franklin's research and her personality. Although sometimes altering history for dramatic effect, the play nevertheless illuminates many of the key issues of how science was and is conducted. ''False Assumptions'' by Lawrence Aronovitch is a play about the life of
Marie Curie Marie Salomea Skłodowska–Curie ( , , ; born Maria Salomea Skłodowska, ; 7 November 1867 – 4 July 1934) was a Polish and naturalized-French physicist and chemist who conducted pioneering research on radioactivity. She was the fir ...
in which Franklin is portrayed as frustrated and angry at the lack of recognition for her scientific contributions. Hostility between the two is also depicted in season 3 of ''Harvey Girls Forever''. Franklin is fictionalised in Marie Benedict's novel ''Her Hidden Genius'', released in January 2022. Franklin was noted as "the chemist that actually discovered DNA" in episode three of the 2019 Netflix series Daybreak (2019 TV series), ''Daybreak''.


Publications

Rosalind Franklin's most notable publications are listed below. The last two were published posthumously. * * * * * * * * * * * * * * * * * * * * * * * * * *


See also

* Timeline of women in science * Cecilia Payne-Gaposchkin, astronomer who discovered the most elemental composition of stars


References


Citations


Sources

* * * * * Elkin, L. O.,
Rosalind Franklin and the Double Helix
Physics Today'' March 2003, pp. 42–48. * * * * * * Holt, J. (2002)
Photo Finish: Rosalind Franklin and the great DNA race
''The New Yorker'' October * * * * * * * * * Watson, J. Letter to ''Science (journal), Science'', ''164'', p. 1539, 27 (1969). * ** *


Further reading

* * * * * * * * * * * * * * * * *


External links

* * * * Recordings by
Aaron Klug Sir Aaron Klug (11 August 1926 – 20 November 2018) was a British biophysicist and chemist. He was a winner of the 1982 Nobel Prize in Chemistry for his development of crystallographic electron microscopy and his structural elucidation of bi ...
at ''Web of Stories'': ** ** ** ** * * * * by Sir Aaron Klug * The first American newspaper coverage of the discovery of the structure of DNA. * * Website for television program first broadcast in 2003 * * Documents from the Churchill Archives Centre, Cambridge. Also available at * * * * * {{DEFAULTSORT:Franklin, Rosalind Elsie 1920 births 1958 deaths 20th-century British biologists 20th-century British chemists 20th-century English scientists 20th-century British women scientists Academics of Birkbeck, University of London Academics of King's College London Alumni of Newnham College, Cambridge Burials at Willesden Jewish Cemetery Carbon scientists Civil Defence Service personnel British crystallographers Deaths from cancer in England Deaths from ovarian cancer English agnostics English Jews English molecular biologists English physical chemists English women biologists English women chemists Franklin family (Anglo-Jewish), Rosalind Jewish agnostics Jewish biophysicists Jewish British scientists Jewish women scientists People educated at Norland Place School People educated at St Paul's Girls' School People from Notting Hill