Editing Frederick Sanger

{{Short description|British biochemist (1918–2013)}}
{{Use British English|date=November 2013}}
{{Use dmy dates|date=August 2022}}
{{Infobox scientist
| image             = Frederick Sanger2.jpg
| honorific_suffix  = {{Postnominals|country=AUS|size=100|OM|CH|CBE|FRS|FAA}}
| birth_date        = {{Birth date |1918|8|13|df=y}}
| death_date        = {{Death date and age|2013|11|19|1918|08|13|df=yes}}
| birth_place       = [[Rendcomb]], Gloucestershire, England
| death_place       = [[Cambridge]], England<ref>{{Cite journal | doi = 10.1038/503442a | title = Seven days: 22–28 November 2013 | journal = Nature | volume = 503 | issue = 7477 | pages = 442–443 | year = 2013 |bibcode = 2013Natur.503..442. | doi-access = free }}</ref>
| field             = [[Biochemistry]]
| work_institutions = {{Plainlist|
* [[University of Cambridge]]
* [[Laboratory of Molecular Biology]]}}
| education         = [[St John's College, Cambridge]] ([[B. A.|BA]], [[PhD]])
| doctoral_advisor  = [[Albert Neuberger]]<ref>{{Cite journal | last1=Allen | first1=A.K. | last2=Muir | first2=H.M. | author2-link=Helen Muir | year=2001 | title=Albert Neuberger. 15 April 1908 – 14 August 1996 | journal=Biographical Memoirs of Fellows of the Royal Society | volume=47 | pages=369–382 | doi=10.1098/rsbm.2001.0021 | jstor=770373 | pmid=15124648| s2cid=72943723 }}</ref>
| doctoral_students = {{Plainlist|
* [[George Brownlee]]<ref name=rsbm>{{Cite journal | last=Brownlee | first=George G. | author-link=George Brownlee | year=2015 | title=Frederick Sanger CBE CH OM. 13 August 1918 – 19 November 2013 | journal=Biographical Memoirs of Fellows of the Royal Society | volume=61 | doi=10.1098/rsbm.2015.0013 | pages=437–466| doi-access=free}}</ref>
* [[Elizabeth Blackburn]]
* [[Rodney Robert Porter|Rodney Porter]]}}
| known_for         = {{Indented plainlist|
* Determining the [[amino acid sequence]] of [[insulin]]
* [[Sanger sequencing]]
* [[Sanger Centre]]}}
| prizes            = {{Plainlist|
* [[Nobel Prize in Chemistry]] (1958, 1980)
* [[Royal Medal]] (1969)
* [[Gairdner Foundation International Award]] (1971)
* [[William Bate Hardy Prize]] (1976)
* [[Copley Medal]] (1977)
* [[Louisa Gross Horwitz Prize]] (1979)}}
| footnotes         = 
| spouse            = Margaret Joan Howe<ref>{{cite web | url=https://www.nobelprize.org/prizes/chemistry/1958/sanger/biographical/ | title=The Nobel Prize in Chemistry 1958 }}</ref>
| relatives         = [[Ruth Sanger]] (first cousin)
| thesis_title      = The metabolism of the amino acid lysine in the animal body
| thesis_url        = http://idiscover.lib.cam.ac.uk/primo-explore/fulldisplay?docid=44CAM_ALMA21428208290003606&context=L&vid=44CAM_PROD&search_scope=default_scope&tab=default_tab&lang=en_US
| thesis_year       = 1943
}}
'''Frederick Sanger''' {{Postnominals|country=AUS|OM|CH|CBE|FRS|FAA}} ({{IPAc-en|ˈ|s|æ|ŋ|ər}}; 13 August 1918&nbsp;– 19 November 2013) was a British [[biochemist]] who received the [[Nobel Prize in Chemistry]] twice.

He won the 1958 Chemistry Prize for determining the [[amino acid]] sequence of [[insulin]] and numerous other proteins, demonstrating in the process that each had a unique, definite structure; this was a foundational discovery for the [[central dogma of molecular biology]].

At the newly constructed [[Laboratory of Molecular Biology]] in Cambridge, he developed and subsequently refined the [[Sanger sequencing|first-ever DNA sequencing technique]], which vastly expanded the number of feasible experiments in molecular biology and remains in widespread use today. The breakthrough earned him the 1980 Nobel Prize in Chemistry, which he shared with [[Walter Gilbert]] and [[Paul Berg]].

He is one of only three people to have won multiple Nobel Prizes in the same category (the others being [[John Bardeen]] in physics and [[Karl Barry Sharpless]] in chemistry),<ref name=nobelfacts>{{Cite web | title=Nobel Prize Facts | publisher=Nobelprize.org | url=https://www.nobelprize.org/nobel_prizes/facts/ | access-date=1 September 2015}}</ref> and one of five persons with [[:Category:Nobel laureates with multiple Nobel awards|two Nobel Prizes]].

==Early life and education==
Frederick Sanger was born on 13 August 1918 in [[Rendcomb]], a small village in [[Gloucestershire]], England, the second son of Frederick Sanger, a [[general practitioner]], and his wife, Cicely Sanger (née Crewdson).<ref name=nobelbio>{{Cite web | title=The Nobel Prize in Chemistry 1958: Frederick Sanger – biography | url= https://www.nobelprize.org/prizes/chemistry/1958/sanger/biographical/ | publisher=Nobelprize.org | access-date=10 August 2020}}</ref> He was one of three children. His brother, Theodore, was only a year older, while his sister May (Mary) was five years younger.<ref name=brownlee>{{Cite web | title=A Life of Research on the Sequences of Proteins and Nucleic Acids: Fred Sanger in conversation with George Brownlee | date=9 October 1992 | url=http://jiscmediahub.ac.uk/record/display/012-00002584 | publisher=Biochemical Society, Edina – Film & Sound Online | access-date=29 April 2013 | archive-url=https://web.archive.org/web/20140313164607/http://jiscmediahub.ac.uk/record/display/012-00002584 | archive-date=13 March 2014 | url-status=dead }}. Subscription required. A 200&nbsp;min interview divided into 44 segments. Notes give the content of each segment. {{Dead link|date=March 2019}}</ref>  His father had worked as an Anglican medical missionary in China but returned to England because of ill health. He was 40 in 1916 when he married Cicely, who was four years younger. Sanger's father converted to [[Quakerism]] soon after his two sons were born and brought up the children as Quakers. Sanger's mother was the daughter of an affluent cotton manufacturer and had a Quaker background, but was not a Quaker.<ref name=brownlee/><!-- This unreliable source http://pennyghael.org.uk/Wilson3.pdf provides genealogical info which is consistent with statements made by Sanger in interviews.
Mother 27 January 1880 – 3 February 1938 – aged 58
Father 20 July 1876 – 6 May 1937 – aged 60
They married in 27 September 1916 -->

When Sanger was around five years old the family moved to the small village of [[Tanworth-in-Arden]] in Warwickshire. The family was reasonably wealthy and employed a governess to teach the children. In 1927, at the age of nine, he was sent to the [[The Downs School (Herefordshire)|Downs School]], a residential preparatory school run by Quakers near [[Malvern, Worcestershire|Malvern]]. His brother Theo was a year ahead of him at the same school. In 1932, at the age of 14, he was sent to the recently established [[Bryanston School]] in Dorset. This used the [[Dalton Plan|Dalton]] system and had a more liberal regime which Sanger much preferred. At the school he liked his teachers and particularly enjoyed scientific subjects.<ref name=brownlee/> Able to complete his [[School Certificate (UK)|School Certificate]] a year early, for which he was awarded seven credits, Sanger was able to spend most of his last year of school experimenting in the laboratory alongside his chemistry master, Geoffrey Ordish, who had originally studied at Cambridge University and been a researcher in the [[Cavendish Laboratory]]. Working with Ordish made a refreshing change from sitting and studying books and awakened Sanger's desire to pursue a scientific career.<ref name=marksearly>{{Cite web | last=Marks | first=Lara | title=Sanger's early life: From the cradle to the laboratory | url=http://www.whatisbiotechnology.org/exhibitions/sanger/early | publisher=What is Biotechnology  | work=The path to DNA sequencing: The life and work of Fred Sanger | access-date=1 September 2015}}</ref> In 1935, prior to heading off to college, Sanger was sent to [[Schule Schloss Salem]] in southern Germany on an exchange program. The school placed a heavy emphasis on athletics, which caused Sanger to be much further ahead in the course material compared to the other students. He was shocked to learn that each day was started with readings from Hitler's [[Mein Kampf]], followed by a [[Sieg Heil]] salute.<ref>{{Cite book |last1=Jeffers |first1=Joe S. |title=Frederick Sanger Two-Time Nobel Laureate in Chemistry |date=2017 |publisher=Springer International| isbn=978-3-319-54707-7 }}</ref>

In 1936 Sanger went to [[St John's College, Cambridge]], to study natural sciences. His father had attended the same college. For Part&nbsp;I of his [[Tripos]] he took courses in physics, chemistry, biochemistry and mathematics but struggled with physics and mathematics.  Many of the other students had studied more mathematics at school. In his second year he replaced physics with physiology. He took three years to obtain his Part&nbsp;I. For his Part&nbsp;II he studied biochemistry and obtained a 1st Class Honours. Biochemistry was a relatively new department founded by [[Gowland Hopkins]] with enthusiastic lecturers who included [[Malcolm Dixon (biochemist)|Malcolm Dixon]], [[Joseph Needham]] and [[Ernest Baldwin]].<ref name=brownlee/>

Both his parents died from cancer during his first two years at Cambridge. His father was 60 and his mother was 58. As an undergraduate Sanger's beliefs were strongly influenced by his Quaker upbringing. He was a pacifist and a member of the [[Peace Pledge Union]]. It was through his involvement with the [[Cambridge Scientists' Anti-War Group]] that he met his future wife, Joan Howe, who was studying economics at [[Newnham College]]. They courted while he was studying for his Part&nbsp;II exams and married after he had graduated in December 1940. Sanger, although brought up and influenced by his religious upbringing, later began to lose sight of his Quaker related ways. He began to see the world through a more scientific lens, and with the growth of his research and scientific development he slowly drifted farther from the faith he grew up with. He had nothing but respect for the religious and states he took two things from it, truth and respect for all life.<ref>{{Cite web|url=https://www.nobelprize.org/prizes/chemistry/1980/sanger/25898-interview-transcript-1980-2/|title=The Nobel Prize in Chemistry 1980}}</ref> Under the [[Military Training Act 1939]] he was provisionally registered as a [[conscientious objector]], and again under the [[National Service (Armed Forces) Act 1939]], before being granted unconditional exemption from military service by a tribunal. In the meantime he undertook training in social relief work at the Quaker centre, Spicelands, Devon and served briefly as a hospital orderly.<ref name=brownlee/>

Sanger began studying for a [[PhD]] in October 1940 under [[Norman Pirie|N.W. "Bill" Pirie]]. His project was to investigate whether edible protein could be obtained from grass. After little more than a month Pirie left the department and [[Albert Neuberger]] became his adviser.<ref name=brownlee/> Sanger changed his research project to study the metabolism of [[lysine]]<ref name="sangerphd">{{Cite thesis |degree=PhD |first=Frederick |last=Sanger |title=The metabolism of the amino acid lysine in the animal body |publisher=University of Cambridge |date=1944 |url=http://ulmss-newton.lib.cam.ac.uk/vwebv/holdingsInfo?bibId=35253 |author-link=Frederick Sanger |access-date=5 December 2013 |archive-date=17 April 2021 |archive-url=https://web.archive.org/web/20210417201357/http://ulmss-newton.lib.cam.ac.uk/vwebv/holdingsInfo?bibId=35253 |url-status=dead }}</ref> and a more practical problem concerning the nitrogen of potatoes.<ref>{{Harvnb|Neuberger|Sanger|1942}}; {{Harvnb|Neuberger|Sanger|1944}}</ref> His thesis had the title, "The metabolism of the amino acid lysine in the animal body". He was examined by [[Charles Harington (chemist)|Charles Harington]] and [[Albert Charles Chibnall]] and awarded his doctorate in 1943.<ref name=brownlee/>

==Research and career==
[[File:Insulin seq vertical.svg|right|thumb|Amino acid sequence of bovine insulin, with [[Disulfide#Occurrence in biology|disulfide bridges]] shown in red]]

===Sequencing insulin===
Neuberger moved to the [[National Institute for Medical Research]] in London, but Sanger stayed in Cambridge and in 1943 joined the group of [[Charles Chibnall]], a protein chemist who had recently taken up the chair in the Department of Biochemistry.<ref>{{Cite web|title= Frederick Sanger, Ph.D. Biography and Interview |website=www.achievement.org|publisher=[[American Academy of Achievement]]|url=https://achievement.org/achiever/frederick-sanger-ph-d/#interview}}</ref> Chibnall had already done some work on the amino acid composition of bovine [[insulin]]<ref>{{Cite journal | last1 = Chibnall | first1 = A. C. | title = Bakerian Lecture: Amino-Acid Analysis and the Structure of Proteins | doi = 10.1098/rspb.1942.0021 | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 131 | issue = 863 | pages = 136–160 | year = 1942 |bibcode = 1942RSPSB.131..136C | s2cid = 85124201 | url = http://rspa.royalsocietypublishing.org/content/royprsa/181/985/210.full.pdf | doi-access = free }} Section on insulin starts on page 153.</ref> and suggested that Sanger look at the amino groups in the protein. Insulin could be purchased from the [[pharmacy]] chain [[Boots UK|Boots]] and was one of the very few proteins that were available in a pure form. Up to this time Sanger had been funding himself. In Chibnall's group he was initially supported by the [[Medical Research Council (UK)|Medical Research Council]] and then from 1944 until 1951 by a [[Beit Memorial Fellowship for Medical Research]].<ref name=nobelbio/>

Sanger's first triumph was to determine the complete [[amino acid]] [[protein sequence|sequence]] of the two polypeptide chains of bovine insulin, A and B, in 1952 and 1951, respectively.<ref>{{Harvnb|Sanger|Tuppy|1951a}}; {{Harvnb|Sanger|Tuppy|1951b}}; {{Harvnb|Sanger|Thompson|1953a}}; {{Harvnb|Sanger|Thompson|1953b}}</ref><ref name=lecture58>{{Citation | last=Sanger | first=F. | year=1958 | title=Nobel lecture: The chemistry of insulin | publisher=Nobelprize.org | url=http://nobelprize.org/nobel_prizes/chemistry/laureates/1958/sanger-lecture.pdf | access-date=18 October 2010 }}. Sanger's Nobel lecture was also published in Science: {{Harvnb|Sanger|1959}}</ref> Prior to this it was widely assumed that proteins were somewhat amorphous. In determining these sequences, Sanger proved that [[protein]]s have a defined chemical composition.<ref name=brownlee/><!--Segment 20-->

To get to this point, Sanger refined a partition chromatography method first developed by [[Richard Laurence Millington Synge]] and [[Archer John Porter Martin]] to determine the composition of amino acids in wool. Sanger used a chemical reagent [[1-fluoro-2,4-dinitrobenzene]] (now, also known as [[Sanger's reagent]], fluorodinitrobenzene, FDNB or DNFB), sourced from [[poisonous gas]] research by Bernard Charles Saunders at the Chemistry Department at Cambridge University. Sanger's reagent proved effective at labelling the  N-terminal amino group at one end of the polypeptide chain.<ref name=marksinsulin>{{Cite web | last=Marks | first=Lara | title=Sequencing proteins: Insulin | url=http://www.whatisbiotechnology.org/exhibitions/sanger/insulin | publisher=What is Biotechnology  | work=The path to DNA sequencing: The life and work of Fred Sanger | access-date=1 September 2015}}</ref> He then partially hydrolysed the insulin into short peptides, either with hydrochloric acid or using an enzyme such as [[trypsin]]. The mixture of peptides was fractionated in two dimensions on a sheet of filter paper, first by [[protein electrophoresis|electrophoresis]] in one dimension and then, perpendicular to that, by [[chromatography]] in the other. The different peptide fragments of insulin, detected with [[ninhydrin]], moved to different positions on the paper, creating a distinct pattern that Sanger called "fingerprints".  The peptide from the N-terminus could be recognised by the yellow colour imparted by the FDNB label and the identity of the labelled amino acid at the end of the peptide determined by complete acid hydrolysis and discovering which dinitrophenyl-amino acid was there.<ref name=brownlee/>

By repeating this type of procedure Sanger was able to determine the sequences of the many peptides generated using different methods for the initial partial hydrolysis. These could then be assembled into the longer [[amino acid sequence|sequence]]s to deduce the complete structure of insulin.  Finally, because the A and B chains are physiologically inactive without the three linking [[disulfide bond]]s (two interchain, one intrachain on A), Sanger and coworkers determined their assignments in 1955.{{sfn|Ryle|Sanger|Smith|Kitai|1955}}<ref name=stretton2002>{{Cite journal | last1 = Stretton | first1 = A.O. | title = The first sequence. Fred Sanger and insulin | journal=Genetics | volume=162 | issue=2 | pages=527–532 | year=2002 | doi = 10.1093/genetics/162.2.527 | pmid=12399368 | pmc=1462286}}</ref> Sanger's principal conclusion was that the two polypeptide chains of the protein insulin had precise amino acid sequences and, by extension, that every protein had a unique sequence.  It was this achievement that earned him his first [[Nobel prize in Chemistry]] in 1958.<ref name=nobel1958>{{Cite web | title=The Nobel Prize in Chemistry 1958: Frederick Sanger| url= http://nobelprize.org/nobel_prizes/chemistry/laureates/1958/ | publisher=Nobelprize.org | access-date=8 October 2010}}</ref> This discovery was crucial to the later [[sequence hypothesis]] of [[Francis Crick]] for developing ideas of how DNA codes for proteins.<ref name=markspath>{{Cite web | last=Marks | first=Lara | title=The path to sequencing nucleic acids | url=http://www.whatisbiotechnology.org/exhibitions/sanger/path | publisher=What is Biotechnology  | work=The path to DNA sequencing: The life and work of Fred Sanger | access-date=1 September 2015}}</ref>

===Sequencing RNA===
From 1951 Sanger was a member of the external staff of the [[Medical Research Council (UK)|Medical Research Council]]<ref name=nobelbio/> and when they opened the [[Laboratory of Molecular Biology]] in 1962, he moved from his laboratories in the Biochemistry Department of the university to the top floor of the new building. He became head of the Protein Chemistry division.<ref name=brownlee/><!--segment 26-->

Prior to his move, Sanger began exploring the possibility of sequencing RNA molecules and began developing methods for separating ribonucleotide fragments generated with specific nucleases. This work he did while trying to refine the sequencing techniques he had developed during his work on insulin.<ref name=markspath/>

The key challenge in the work was finding a pure piece of RNA to sequence. In the course of the work he discovered in 1964, with Kjeld Marcker, the [[formylmethionine|formylmethionine tRNA]] which initiates protein synthesis in bacteria.<ref>{{Harvnb|Marcker|Sanger|1964}}</ref> He was beaten in the race to be the first to sequence a [[tRNA]] molecule by a group led by [[Robert W. Holley|Robert Holley]] from [[Cornell University]], who published the sequence of the 77 ribonucleotides of [[alanine]] tRNA from ''[[Saccharomyces cerevisiae]]'' in 1965.<ref>{{Cite journal | last1 = Holley | first1 = R. W. | last2 = Apgar | first2 = J. | last3 = Everett | first3 = G. A. | last4 = Madison | first4 = J. T. | last5 = Marquisee | first5 = M. | last6 = Merrill | first6 = S. H. | last7 = Penswick | first7 = J. R. | last8 = Zamir | first8 = A. | doi = 10.1126/science.147.3664.1462 | title = Structure of a Ribonucleic Acid | journal = Science | volume = 147 | issue = 3664 | pages = 1462–1465 | year = 1965 | pmid =  14263761|bibcode = 1965Sci...147.1462H | s2cid = 40989800 }}</ref> By 1967 Sanger's group had determined the nucleotide sequence of the [[5S ribosomal RNA]] from ''[[Escherichia coli]]'', a small RNA of 120 nucleotides.<ref>{{Harvnb|Brownlee|Sanger|Barrell|1967}}; {{Harvnb|Brownlee|Sanger|Barrell|1968}}</ref>

===Sequencing DNA===
{{main|DNA sequencing}}
Sanger then turned to sequencing DNA, which would require an entirely different approach. He looked at different ways of using [[DNA polymerase I]] from ''E. coli'' to copy single-stranded DNA.<ref>{{Harvnb|Sanger|Donelson|Coulson|Kössel|1973}}</ref> In 1975, together with [[Alan Coulson]], he published a sequencing procedure using DNA polymerase with radiolabelled nucleotides that he called the "Plus and Minus" technique.<ref>{{Harvnb|Sanger|Coulson|1975}}</ref><ref name=lecture80>{{Cite web| last=Sanger | first=F. | year=1980 | title=Nobel lecture: Determination of nucleotide sequences in DNA | publisher=Nobelprize.org | url=https://www.nobelprize.org/uploads/2018/06/sanger-lecture-1.pdf | access-date=15 September 2019 }}</ref> This involved two closely related methods that generated short oligonucleotides with defined 3' termini. These could be fractionated by [[electrophoresis]] on a [[polyacrylamide]] gel and visualised using [[autoradiography]]. The procedure could sequence up to 80 nucleotides in one go and was a big improvement on what had gone before, but was still very laborious. Nevertheless, his group were able to sequence most of the 5,386 nucleotides of the single-stranded [[bacteriophage]] [[Phi X 174|φX174]].<ref>{{Harvnb|Sanger|Air|Barrell|Brown|1977}}</ref> This was the first fully sequenced DNA-based genome. To their surprise they discovered that the [[coding region]]s of some of the genes overlapped with one another.<ref name=rsbm/>

In 1977 Sanger and colleagues introduced the "dideoxy" chain-termination method for sequencing DNA molecules, also known as the "[[Sanger method]]".<ref name=lecture80/>{{sfn|Sanger|Nicklen|Coulson|1977}} This was a major breakthrough and allowed long stretches of DNA to be rapidly and accurately sequenced. It earned him his second Nobel prize in Chemistry in 1980, which he shared with [[Walter Gilbert]] and [[Paul Berg]].<ref name=nobel1980>{{Cite web | title=The Nobel Prize in Chemistry 1980: Paul Berg, Walter Gilbert, Frederick Sanger | url=http://nobelprize.org/nobel_prizes/chemistry/laureates/1980/ | publisher=Nobelprize.org | access-date=8 October 2010}}</ref> The new method was used by Sanger and colleagues to sequence human mitochondrial DNA (16,569 base pairs)<ref>{{Harvnb|Anderson|Bankier|Barrell|De Bruijn|1981}}</ref> and bacteriophage λ (48,502 base pairs).<ref>{{Harvnb|Sanger|Coulson|Hong| Hill|1982}}</ref> The dideoxy method was eventually used to sequence the entire [[human genome]].<ref name="obituarynature">{{Cite journal | last1 = Walker | first1 = John| author-link = John E. Walker| title =Frederick Sanger (1918–2013) Double Nobel-prizewinning genomics pioneer| doi = 10.1038/505027a | journal = Nature | volume = 505 | issue = 7481 | pages = 27 | year = 2014 | pmid =  24380948|bibcode = 2014Natur.505...27W | doi-access = free }}</ref>

===Postgraduate students===
During the course of his career Sanger supervised more than ten PhD students, two of whom went on to also win Nobel Prizes. His first graduate student was [[Rodney Robert Porter|Rodney Porter]] who joined the research group in 1947.<ref name=rsbm/> Porter later shared the 1972 [[Nobel Prize in Physiology or Medicine]] with [[Gerald Edelman]] for his work on the chemical structure of [[antibodies]].<ref>{{ cite web | title=The Nobel Prize in Physiology or Medicine 1972 | publisher=Nobelprize.org | url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/1972/ | access-date=1 September 2015 }}</ref> [[Elizabeth Blackburn]] studied for a PhD in Sanger's laboratory between 1971 and 1974.<ref name=rsbm/><ref name=blackphd>{{Cite thesis|degree=PhD|publisher=University of Cambridge|title=Sequence studies on bacteriophage ØX174 DNA by transcription|first=E. H.|last=Blackburn|date=1974|url=http://ethos.bl.uk/OrderDetails.do?uin=uk.bl.ethos.449954|access-date=4 July 2015|archive-date=5 March 2016|archive-url=https://web.archive.org/web/20160305133053/http://ethos.bl.uk/OrderDetails.do?uin=uk.bl.ethos.449954|url-status=dead}}</ref> She shared the 2009 Nobel Prize in Physiology or Medicine with [[Carol W. Greider]] and [[Jack W. Szostak]] for her work on [[telomere]]s and the action of [[telomerase]].<ref>{{ cite web | title=The Nobel Prize in Physiology or Medicine 2009 | publisher=Nobelprize.org | url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/2009/ | access-date=1 September 2015 }}</ref>

===Sanger's rule===
{{blockquote|... anytime you get technical development that’s two to threefold or more efficient, accurate, cheaper, a whole range of experiments opens up.<ref>{{cite web|title=Schlessinger, David|date=March 2018|website=National Human Genome Research Institute (NHGRI, genome.gov)|url=https://www.genome.gov/sites/default/files/media/files/2019-05/david_schlessinger_transcript.pdf}}</ref>}}

This rule should not be confused with Terence Sanger's rule, which is related to [[Oja's rule]].

===Awards and honours===
{{As of|2015}}, Sanger is one of the only two people to have been awarded the Nobel Prize in Chemistry twice (the other being [[Karl Barry Sharpless]] in 2001 and 2022'')'', and one of only five two-time Nobel laureates: The other four were [[Marie Curie]] ([[Nobel Prize in Physics|Physics]], 1903 and [[Nobel Prize in Chemistry|Chemistry]], 1911), [[Linus Pauling]] ([[Nobel Prize in Chemistry|Chemistry]], 1954 and [[Nobel Peace Prize|Peace]], 1962), [[John Bardeen]] (twice [[Nobel Prize in Physics|Physics]], 1956 and 1972), and [[Karl Barry Sharpless]] (twice [[Nobel Prize in Chemistry|Chemistry]], 2001 and 2022).<ref name=nobelfacts/>

* Elected [[List of Fellows of the Royal Society elected in 1954|Fellow of the Royal Society (FRS) in 1954]]<ref name=rsbm/>
* [[Commander of the Order of the British Empire]] (CBE) – 1963<ref name=rsbm/>
* [[Member of the Order of the Companions of Honour]] (CH) – 1981<ref name=rsbm/>
* [[Member of the Order of Merit]] (OM) – 1986<ref name=rsbm/>
* Corresponding Member of the [[Australian Academy of Science]] – 1982<ref name=rsbm/>
* [[William Bate Hardy Prize]] – 1976<ref name=rsbm/>
* [[Nobel Prize in Chemistry]] – 1958, 1980<ref name=nobel1958/><ref name=nobel1980/>
* [[Corday–Morgan Medal]] – 1951<ref name=rsbm/>
* [[Royal Medal]] – 1969<ref name=rsbm/>
* [[Gairdner Foundation International Award]] – 1971<ref name=rsbm/>
* [[Copley Medal]] – 1977<ref name=rsbm/>
* G.W. Wheland Award – 1978<ref name=rsbm/>
* [[Louisa Gross Horwitz Prize]] of [[Columbia University]] – 1979<ref name=rsbm/>
* [[Albert Lasker Award for Basic Medical Research]] – 1979<ref name=rsbm/>
* [[Association of Biomolecular Resource Facilities]] Award – 1994<ref>{{Cite web | title=The ABRF Award for Outstanding Contributions to Biomolecular Technologies | publisher=Association of Biomolecular Resource facilities | url=https://abrf.org/awards/abrf-award-outstanding-contributions-biomolecular-technologies | access-date=11 August 2020 | archive-date=6 August 2020 | archive-url=https://web.archive.org/web/20200806131051/https://abrf.org/awards/abrf-award-outstanding-contributions-biomolecular-technologies | url-status=dead }}</ref>
* Golden Plate Award of the [[Academy of Achievement|American Academy of Achievement]] – 2000<ref>{{Cite web|title= Golden Plate Awardees of the American Academy of Achievement |website=www.achievement.org|publisher=[[American Academy of Achievement]]|url=  https://www.achievement.org/our-history/golden-plate-awards/#science-exploration/}}</ref><ref>{{Cite news |title= Summit Overview Photo| url=  https://achievement.org/summit/|quote= Awards Council member and Nobel Prize laureate Dr. Charles H. Townes presenting the American Academy of Achievement’s Golden Plate Award to British biochemist Dr. Frederick Sanger, recipient of two Nobel Prizes in Chemistry, at the 2000 Summit in Hampton Court.}}</ref>
* Citation for Chemical Breakthrough Award from the Division of History of Chemistry of the American Chemical Society – 2016<ref name="Award">{{Cite web|title=2016 Awardees|url=http://www.scs.illinois.edu/~mainzv/HIST/awards/CCB-2016_Awardees.php|website=American Chemical Society, Division of the History of Chemistry|publisher=University of Illinois at Urbana-Champaign School of Chemical Sciences|date=2016|access-date=14 June 2017|archive-date=6 May 2017|archive-url=https://web.archive.org/web/20170506160732/http://www.scs.illinois.edu/~mainzv/HIST/awards/CCB-2016_Awardees.php|url-status=dead}}</ref><ref name="Breakthrough">{{Cite web|title=Citation for Chemical Breakthrough Award|url=http://www.scs.illinois.edu/~mainzv/HIST/awards/Citations/2016-ACS%20Rendering_Sanger.pdf|archive-url=https://web.archive.org/web/20170506173200/http://www.scs.illinois.edu/~mainzv/HIST/awards/Citations/2016-ACS%20Rendering_Sanger.pdf|url-status=dead|archive-date=6 May 2017|website=American Chemical Society, Division of the History of Chemistry|publisher=University of Illinois at Urbana-Champaign School of Chemical Sciences|date=2016|access-date=14 June 2017}}</ref>{{sfn|Sanger|Nicklen|Coulson|1977}}

The [[Wellcome Trust Sanger Institute]] (formerly the ''Sanger Centre'') is named in his honour.<ref name=rsbm/>

==Personal life==

===Marriage and family===
Sanger married Margaret Joan Howe (not to be confused with [[Margaret Sanger]], the American pioneer of birth control) in 1940. She died in 2012. They had three children—Robin, born in 1943, Peter born in 1946 and Sally Joan born in 1960.<ref name=nobelbio/> He said that his wife had "contributed more to his work than anyone else by providing a peaceful and happy home."<ref name=Telegraph_obit/>

===Later life===
[[File:EBI and Sanger Center, Genome campus, Cambridgeshire.jpg|thumb|right|300px| The Sanger Institute]]
Sanger retired in 1983, aged 65, to his home, "Far Leys", in [[Swaffham Bulbeck]] outside Cambridge.<ref name=rsbm/>

In 1992, the [[Wellcome Trust]] and the Medical Research Council founded the Sanger Centre (now the [[Sanger Institute]]), named after him.<ref name="Our people">{{Cite web | title=Frederick Sanger | url=http://www.sanger.ac.uk/about/people/biographies/fsanger.html | publisher=Wellcome Trust Sanger Institute | access-date=12 October 2010 | url-status=dead | archive-url=https://web.archive.org/web/20110407064631/http://www.sanger.ac.uk/about/people/biographies/fsanger.html | archive-date=7 April 2011 | df=dmy-all }}</ref> The institute is on the [[Wellcome Trust Genome Campus]] near [[Hinxton]], only a few miles from Sanger's home. He agreed to having the Centre named after him when asked by [[John Sulston]], the founding director, but warned, "It had better be good."<ref name="Our people"/> It was opened by Sanger in person on 4 October 1993, with a staff of fewer than 50 people, and went on to take a leading role in the [[Human Genome Project|sequencing of the human genome]].<ref name="Our people"/> The Institute had about 900 people in 2020 and is one of the world's largest [[genomic]] research centres.

Sanger said he found no evidence for a God so he became an agnostic.<ref name=hargittai>{{Cite journal | last=Hargittai | first=István | title=Interview: Frederick Sanger | journal=The Chemical Intelligencer | volume=4 | issue=2 | pages=6–11 | publisher=Springer-Verlag | place=New York | date=April 1999 }}. This interview, which took place on 16 September 1997, was republished in {{Cite book | last=Hargittai | first=István | year=2002 | title=Candid science II: conversations with famous biomedical scientists | chapter=Chapter 5: Frederick Sanger | publisher=Imperial College Press | place=London | isbn=978-1-86094-288-4 | pages=73–83}}</ref> In an interview published in the ''[[The Times|Times]]'' newspaper in 2000 Sanger is quoted as saying: "My father was a committed Quaker and I was brought up as a Quaker, and for them truth is very important. I drifted away from those beliefs – one is obviously looking for truth, but one needs some evidence for it. Even if I wanted to believe in God I would find it very difficult. I would need to see proof."<ref name=times>{{Cite web |last=Ahuja |first=Anjana |title=The double Nobel laureate who began the book of life |newspaper=[[The Times]] |location=London |date=12 January 2000 |page=40 |url=http://www.warwick.ac.uk/fac/sci/Chemistry/MedChem/MedChemInfo/info/Sanger.html |archive-url=https://web.archive.org/web/20081211053639/http://www.warwick.ac.uk/fac/sci/Chemistry/MedChem/MedChemInfo/info/Sanger.html |archive-date=11 December 2008 |via=warwick.ac.uk |access-date=18 October 2010 |url-status=dead }}</ref>

He declined the offer of a [[Orders, decorations, and medals of the United Kingdom#Knighthood|knighthood]], as he did not wish to be addressed as "Sir". He is quoted as saying, "A knighthood makes you different, doesn't it, and I don't want to be different." In 1986 he accepted admission to the [[Order of Merit (Commonwealth)|Order of Merit]], which can have only 24 living members.<ref name=Telegraph_obit/><ref name=hargittai/><ref name=times/>

In 2007 the British [[Biochemical Society]] was given a grant by the [[Wellcome Trust]] to catalogue and preserve the 35 laboratory notebooks in which Sanger recorded his research from 1944 to 1983. In reporting this matter, ''[[Science (journal)|Science]]'' noted that Sanger, "the most self-effacing person you could hope to meet", was spending his time gardening at his [[Cambridgeshire]] home.<ref>{{Cite journal| title=Newsmakers: A Life in Science | editor-last=Bhattachjee | editor-first=Yudhijit | journal=Science | volume=317 | issue=5840 | page=879 | year=2007 | url=http://www.sciencemag.org/content/vol317/issue5840/newsmakers.dtl?cookietest=yes|doi=10.1126/science.317.5840.879e | s2cid=220092058 }}</ref>

Sanger died in his sleep at [[Addenbrooke's Hospital]] in [[Cambridge]] on 19 November 2013.<ref name=Telegraph_obit>{{Cite news | url=https://www.telegraph.co.uk/news/obituaries/science-obituaries/10462574/Frederick-Sanger-OM.html |archive-url=https://ghostarchive.org/archive/20220112/https://www.telegraph.co.uk/news/obituaries/science-obituaries/10462574/Frederick-Sanger-OM.html |archive-date=12 January 2022 |url-access=subscription |url-status=live | newspaper=The Telegraph | title= Frederick Sanger, OM | access-date= 20 November 2013 | date=20 November 2013 }}{{cbignore}}</ref><ref name=death>{{Cite news|url=https://www.bbc.co.uk/news/science-environment-25020112 |title=Frederick Sanger: Nobel Prize winner dies at 95 |publisher=BBC.co.uk |date=20 November 2013 |access-date=20 November 2013}}</ref> As noted in his obituary, he had described himself as "just a chap who messed about in a lab",<ref>{{Cite news | title= Frederick Sanger: Unassuming British biochemist whose pivotal and far-reaching discoveries made him one of a handful of double Nobel prizewinners | location = London | newspaper=The Times | date=21 November 2013 |page=63 }}</ref> and "academically not brilliant".<ref>{{Cite web |url=https://theconversation.com/frederick-sangers-achievements-cannot-be-overstated-20596 |title=Frederick Sanger's achievements cannot be overstated |work=The Conversation |date=21 November 2013 }}</ref>

== Global policy ==
He was one of the signatories of the agreement to convene a convention for drafting a [[world constitution]].<ref>{{Cite web |title=Letters from Thane Read asking Helen Keller to sign the World Constitution for world peace. 1961 |url=https://www.afb.org/HelenKellerArchive?a=d&d=A-HK01-07-B149-F04-022.1.8 |access-date=2023-07-01 |website=Helen Keller Archive |publisher=American Foundation for the Blind}}</ref><ref>{{Cite web |title=Letter from World Constitution Coordinating Committee to Helen, enclosing current materials |url=https://www.afb.org/HelenKellerArchive?a=d&d=A-HK01-07-B154-F05-028.1.6 |access-date=2023-07-03 |website=Helen Keller Archive |publisher=American Foundation for the Blind}}</ref> As a result, for the first time in human history, a [[World Constituent Assembly]] convened to draft and adopt a [[Constitution for the Federation of Earth]].<ref>{{Cite web |title=Preparing earth constitution {{!}} Global Strategies & Solutions {{!}} The Encyclopedia of World Problems |url=http://encyclopedia.uia.org/en/strategy/193465 |url-status=dead |access-date=2023-07-15 |website=The Encyclopedia of World Problems {{!}} Union of International Associations (UIA) |archive-date=19 July 2023 |archive-url=https://web.archive.org/web/20230719215501/http://encyclopedia.uia.org/en/strategy/193465 }}</ref>

==Selected publications==
{{Refbegin|30em}}
*{{Citation| last1=Neuberger | first1=A. | last2=Sanger | first2=F. | year=1942 | title=The nitrogen of the potato | journal=Biochemical Journal | volume=36| issue=7–9 | pages=662–671 | pmid=16747571 | pmc=1266851 | doi=10.1042/bj0360662 }}.
*{{Citation | last1=Neuberger | first1=A. | last2=Sanger | first2=F. | year=1944 | title= The metabolism of lysine | journal=Biochemical Journal | volume=38 | issue=1 | pages=119–125 | pmid=16747737 | pmc=1258037 | doi=10.1042/bj0380119 }}.
*{{Citation | last=Sanger | first=F. | year=1945 | title=The free amino groups of insulin | journal=Biochemical Journal | volume=39 | issue=5 | pages=507–515 | pmid=16747948 | pmc=1258275 | ref=none | doi=10.1042/bj0390507}}.
*{{Citation | last=Sanger | first=F. | year=1947 | title= Oxidation of insulin by performic acid | journal=Nature | volume=160 | issue=4061 | pages=295–296 | pmid=20344639 | doi=10.1038/160295b0 | ref=none |bibcode = 1947Natur.160..295S | s2cid=4127677 }}.
*{{Citation | last1=Porter | first1=R.R. | last2=Sanger | first2=F. | year=1948 | title=The free amino groups of haemoglobins | journal=Biochemical Journal | volume=42 | issue=2 | pages=287–294 | pmid=16748281 | pmc=1258669 | ref=none| doi=10.1042/bj0420287 }}.
*{{Citation | last=Sanger | first=F. | year=1949a | title=Fractionation of oxidized insulin | journal= Biochemical Journal | volume=44 | issue=1 | pages=126–128 | pmid=16748471  | pmc=1274818 | ref=none | doi=10.1042/bj0440126 }}.
*{{Citation | last=Sanger | first=F. | year=1949b | title=The terminal peptides of insulin | journal= Biochemical Journal | volume=45 | issue=5 | pages=563–574 | pmid=15396627  | pmc=1275055 | ref=none| doi=10.1042/bj0450563 }}.
*{{Citation | last1= Sanger | first1=F. | last2=Tuppy | first2=H. | year=1951a | title= The amino-acid sequence in the phenylalanyl chain of insulin. 1. The identification of lower peptides from partial hydrolysates | journal=Biochemical Journal | volume=49 | issue= 4 | pages=463–481 | pmid=14886310 | pmc=1197535 | doi=10.1042/bj0490463}}.
*{{Citation | last1= Sanger | first1=F. | last2=Tuppy | first2=H. | year=1951b | title= The amino-acid sequence in the phenylalanyl chain of insulin. 2. The investigation of peptides from enzymic hydrolysates | journal=Biochemical Journal | volume=49 | issue= 4 | pages=481–490 | pmid=14886311 | pmc=1197536 | doi=10.1042/bj0490481}}.
*{{Citation | last1= Sanger | first1=F. | last2= Thompson | first2=E.O.P. | year=1953a | title= The amino-acid sequence in the glycyl chain of insulin. 1. The identification of lower peptides from partial hydrolysates | journal=Biochemical Journal | volume=53 | issue= 3 | pages=353–366 | pmid=13032078 | pmc=1198157 | doi=10.1042/bj0530353}}.
*{{Citation | last1= Sanger | first1=F. | last2= Thompson | first2=E.O.P. | year=1953b | title= The amino-acid sequence in the glycyl chain of insulin. 2. The investigation of peptides from enzymic hydrolysates | journal=Biochemical Journal | volume=53 | issue= 3 | pages=366–374 | pmid=13032079 | pmc=1198158 | doi=10.1042/bj0530366}}.
*{{Citation | last1=Sanger | first1=F. | last2=Thompson | first2=E.O.P. | last3=Kitai | first3=R. | year=1955 | title=The amide groups of insulin | journal=Biochemical Journal | volume=59 | issue=3 | pages=509–518 | pmid=14363129 | pmc=1216278 | ref=none | doi=10.1042/bj0590509 }}.
*{{Citation | last1=Ryle | first1=A.P. | last2=Sanger | first2=F. | last3=Smith | first3=L.F. | last4=Kitai | first4=R. | year=1955 | title=The disulphide bonds of insulin | journal=Biochemical Journal | volume=60 | issue=4 | pages=541–556 | pmid=13249947 | pmc=1216151 | doi=10.1042/bj0600541 }}.
*{{Citation | last1=Brown | first1=H. | last2=Sanger | first2=F. | last3=Kitai | first3=R. | year=1955 | title= The structure of pig and sheep insulins | journal=Biochemical Journal | volume=60 | issue=4 | pages=556–565 | pmid=13249948 | pmc=1216152 | ref=none| doi=10.1042/bj0600556 }}.
*{{Citation | last=Sanger | first=F. | year=1959 | title= Chemistry of Insulin: determination of the structure of insulin opens the way to greater understanding of life processes | journal=Science | volume=129 | pages=1340–1344 | doi=10.1126/science.129.3359.1340 | pmid=13658959 | issue=3359|bibcode = 1959Sci...129.1340G }}.
*{{Citation | last1=Milstein | first1=C. | last2= Sanger | first2=F. | year=1961 | title= An amino acid sequence in the active centre of phosphoglucomutase | journal=Biochemical Journal | volume=79 | pages=456–469 | pmid=13771000 | pmc=1205670 | issue=3 | ref=none| doi=10.1042/bj0790456 }}.
*{{Citation | last1=Marcker | first1=K. | last2=Sanger | first2=F. | year=1964 | title=''N''-formyl-methionyl-S-RNA | journal=Journal of Molecular Biology | volume=8 | pages=835–840 | pmid=14187409 | doi=10.1016/S0022-2836(64)80164-9 | issue=6 }}.
*{{Citation | last1=Sanger | first1=F. | last2=Brownlee | first2=G.G. | last3=Barrell | first3=B.G. | year=1965 | title= A two-dimensional fractionation procedure for radioactive nucleotides | journal=Journal of Molecular Biology | volume=13 | issue=2 | pages=373–398 | pmid=5325727 | doi=10.1016/S0022-2836(65)80104-8 | ref=none }}.
*{{Citation | last1=Brownlee | first1=G.G.| last2=Sanger | first2=F. | last3=Barrell | first3=B.G. | year=1967 | title= Nucleotide sequence of 5S-ribosomal RNA from ''Escherichia coli'' | journal=Nature | volume=215 | issue=5102 | pages=735–736 | pmid=4862513 | doi=10.1038/215735a0 |bibcode = 1967Natur.215..735B | s2cid=4270186}}.
*{{Citation | last1=Brownlee | first1=G.G. | last2=Sanger | first2=F. | year=1967 | title= Nucleotide sequences from the low molecular weight ribosomal RNA of ''Escherichia coli'' | journal=Journal of Molecular Biology | volume=23 | issue=3 | pages=337–353 | pmid=4291728 | doi=10.1016/S0022-2836(67)80109-8 | ref=none }}.
*{{Citation | last1=Brownlee | first1=G.G.| last2=Sanger | first2=F. | last3=Barrell | first3=B.G. | year=1968 | title= The sequence of 5S ribosomal ribonucleic acid | journal=Journal of Molecular Biology | volume=34 | issue=3 | pages=379–412 | pmid=4938553 | doi=10.1016/0022-2836(68)90168-X }}.
*{{Citation | last1=Adams | first1=J.M. | last2=Jeppesen | first2=P.G. | last3=Sanger | first3=F. | last4=Barrell | first4=B.G. | year=1969 | title= Nucleotide sequence from the coat protein cistron of R17 bacteriophage RNA | journal=Nature | volume=223 | issue=5210 | pages=1009–1014 | pmid=5811898 | doi=10.1038/2231009a0 |bibcode = 1969Natur.223.1009A | s2cid=4152602 | ref=none}}.
*{{Citation | last1=Barrell | first1=B.G. | last2=Sanger | first2=F. | year=1969 | title=The sequence of phenylalanine tRNA from ''E. coli'' | journal=FEBS Letters | volume=3 | issue=4 | pages=275–278 | pmid=11947028 | doi=10.1016/0014-5793(69)80157-2 | s2cid=34155866 | ref=none| doi-access=free | bibcode=1969FEBSL...3..275B }}.
*{{Citation | last1=Jeppesen | first1=P.G. | last2=Barrell | first2=B.G. | last3=Sanger | first3=F. | last4=Coulson | first4=A.R. | year=1972 | title= Nucleotide sequences of two fragments from the coat-protein cistron of bacteriophage R17 ribonucleic acid | journal=Biochemical Journal | volume=128 | issue=5 | pages=993–1006 | pmid=4566195 | pmc=1173988 | ref=none | doi=10.1042/bj1280993h}}.
*{{Citation | doi=10.1073/pnas.70.4.1209 | last1=Sanger | first1=F. | last2=Donelson | first2=J.E. | last3=Coulson | first3=A.R. | last4=Kössel | first4=H. | last5=Fischer | first5=D. | year=1973 | title= Use of DNA Polymerase I Primed by a Synthetic Oligonucleotide to Determine a Nucleotide Sequence in Phage f1 DNA | journal=Proceedings of the National Academy of Sciences USA | volume=70 | issue=4 | pages=1209–1213 | pmid=4577794 | pmc=433459 |bibcode = 1973PNAS...70.1209S | doi-access=free }}.
*{{Citation | last1=Sanger | first1=F. | last2=Coulson | first2=A.R. | year=1975 | title= A rapid method for determining sequences in DNA by primed synthesis with DNA polymerase | journal=Journal of Molecular Biology | volume=94 | issue=3 | pages=441–448 | pmid=1100841 | doi=10.1016/0022-2836(75)90213-2 }}.
*{{Citation | doi=10.1073/pnas.74.12.5463 | last1=Sanger| first1=F. | last2=Nicklen| first2=S. | last3=Coulson| first3=A.R.|title=DNA sequencing with chain-terminating inhibitors| journal=Proceedings of the National Academy of Sciences USA | year=1977 | volume=74 | issue=12 | pages=5463–5467 | pmid=271968 | pmc=431765 |bibcode = 1977PNAS...74.5463S | doi-access=free}}. According to the [[Institute for Scientific Information]] (ISI) database, by October 2010 this paper had been cited over 64,000 times.
*{{Citation | last1=Sanger | first1=F. | last2=Air | first2=G.M. | last3=Barrell | first3=B.G. | last4=Brown | first4=N.L. | last5=Coulson | first5=A.R. | last6=Fiddes | first6=C.A. | last7=Hutchinson | first7=C.A. | last8=Slocombe | first8=P.M. | last9=Smith | first9=M. | year=1977 | title= Nucleotide sequence of bacteriophage φX174 DNA | journal=Nature | volume=265 | issue=5596 | pages=687–695 | pmid=870828 | doi=10.1038/265687a0 |bibcode = 1977Natur.265..687S  | s2cid=4206886 }}.
*{{Citation | last1=Sanger | first1=F. | last2=Coulson | first2=A.R. | year=1978 | title= The use of thin acrylamide gels for DNA sequencing | journal= FEBS Letters | volume=87 | issue=1 | pages=107–110 | pmid=631324 | doi=10.1016/0014-5793(78)80145-8 | s2cid=1620755 | ref=none| doi-access= | bibcode=1978FEBSL..87..107S }}.
*{{Citation | last1=Sanger |first1=F. | last2=Coulson | first2=A.R. | last3=Barrell | first3=B.G. | last4=Smith | first4= A.J. | last5=Roe | first5=B.A. | year=1980 | title= Cloning in single-stranded bacteriophage as an aid to rapid DNA sequencing | journal=Journal of Molecular Biology | volume=143 | issue=2 | pages=161–178 | pmid=6260957 | doi=10.1016/0022-2836(80)90196-5 | ref=none }}.
*{{Citation | last1=Anderson | first1=S. | year=1981 | last2=Bankier | first2=A.T. | last3=Barrell | first3=B.G. | last4=De Bruijn | first4=M.H. | last5=Coulson | first5=A.R. | last6=Drouin | first6=J. | last7=Eperon | first7=I.C. | last8=Nierlich | first8=D.P. | last9=Roe | first9=B.A. | last10=Sanger | first10=F. | last11=Schreier | first11=P.H. | last12=Smith | first12=A.J. | last13=Staden | first13=R. |last14=Young | first14=I.G. | title= Sequence and organization of the human mitochondrial genome | journal=Nature | volume=290 | issue=5806 | pages=457–465 | pmid=7219534 | doi=10.1038/290457a0 |bibcode = 1981Natur.290..457A  | s2cid=4355527 }}.
*{{Citation | last1=Anderson | first1=S. | last2=De Bruijn | year=1982 | first2=M.H. | last3=Coulson | first3=A.R. | last4=Eperon | first4=I.C. | last5=Sanger | first5=F. | last6=Young | first6=I.G. | title= Complete sequence of bovine mitochondrial DNA. Conserved features of the mammalian mitochondrial genome | journal=Journal of Molecular Biology | volume=156 | issue=4 | pages=683–717 | pmid=7120390 | doi=10.1016/0022-2836(82)90137-1 | ref=none}}.
*{{Citation | last1=Sanger | first1=F. | last2=Coulson | first2=A.R. | last3=Hong | first3=G.F. | last4=Hill | first4=D.F. | last5=Petersen | first5=G.B. |authorlink4=Diana Hill (scientist) |authorlink5=George Petersen (biochemist) | year=1982 | title= Nucleotide sequence of bacteriophage λ DNA | journal=Journal of Molecular Biology | volume=162 | issue=4 | pages=729–773 | pmid=6221115 | doi=10.1016/0022-2836(82)90546-0 }}.
*{{Citation | last=Sanger | first=F. | year=1988 | title=Sequences, sequences, and sequences | journal=Annual Review of Biochemistry | volume=57 | pages=1–28 | pmid=2460023 | doi=10.1146/annurev.bi.57.070188.000245 | ref=none | doi-access=free }}.
{{Refend}}

==References==
=== Citations ===
{{Reflist}}

=== Bibliography ===
{{div col|colwidth=35em}}
*{{Cite book | last=Brownlee | first=George G. | author-link=George Brownlee | year=2014 | title=Fred Sanger, double Nobel laureate: a biography | publisher=Cambridge University Press | place=Cambridge, UK | isbn=978-1-107-08334-9 | ref=none }} Chapters 4-6 contain the 1992 interview that the author conducted with Sanger.
*{{Citation|  last=Finch | first=John | year=2008 | title=A Nobel Fellow on every floor: a history of the Medical Research Council Laboratory of Molecular Biology | publisher=Medical Research Council | place=Cambridge | isbn=978-1-84046-940-0 | ref=none}}.
*{{ cite journal | last=García-Sancho | first=Miguel | year=2010 | title=A new insight into Sanger's development of sequencing: from proteins to DNA, 1943–1977 | journal=Journal of the History of Biology | volume=43 | issue=2 | pages=265–323 | doi=10.1007/s10739-009-9184-1 | pmid=20665230 | s2cid=1134280 | url=https://www.pure.ed.ac.uk/ws/files/16303515/sangerjhb.pdf | ref=none | hdl=20.500.11820/e4febe48-772a-4f47-a1c5-a5ca89505367 | hdl-access=free }}
*{{Citation | last1=Sanger | first1=F. | last2= Dowding | first2=M. | year=1996 | title=Selected Papers of Frederick Sanger: with commentaries | publisher= World Scientific | place=Singapore | isbn= 978-981-02-2430-1 | ref=none }}.
*{{Citation | title=Interviews with Nobel Prize–winning scientists: Dr Frederick Sanger | url=https://www.bbc.co.uk/archive/scientists/10603.shtml | year=c. 1985 | publisher=British Broadcasting Corporation | ref=none }}. Interviewed by [[Lewis Wolpert]]. Duration 1 hour.
{{div col end}}

==External links ==
{{Commons category}}
*[https://www.sanger.ac.uk/ The Sanger Institute]
*[https://www.nobelprize.org/prizes/chemistry/1958/summary/ About the 1958 Nobel Prize]
*[https://www.nobelprize.org/prizes/chemistry/1980/summary/ About the 1980 Nobel Prize]
*[https://web.archive.org/web/20041026014225/http://www.vega.org.uk/series/facetoface/sanger/index.php Fred Sanger] 2001 Video Documentary by The Vega Science Trust
* {{NPG name}}
*[https://www.sms.cam.ac.uk/media/1130236 Frederick Sanger interviewed by Alan Macfarlane, 24 August 2007 (video)], also available on {{YouTube|0JsrvWYS7zY}}. Duration 57 minutes.
*[http://wellcomelibrary.org/using-the-library/subject-guides/genetics/makers-of-modern-genetics/digitised-archives/fred-sanger/ Frederick Sanger archive collection] – Wellcome Library finding aid for the digitised collection.
* {{Nobelprize}}

{{Nobel Prize in Chemistry}}
{{1958 Nobel Prize winners}}
{{1980 Nobel Prize winners}}
{{Copley Medallists 1951–2000}}
{{World Constitutional Convention call signatories}}
{{Portal bar|Biography|Biology|Viruses}}

{{Authority control}}

{{DEFAULTSORT:Sanger, Frederick}}
[[Category:1918 births]]
[[Category:2013 deaths]]
[[Category:Alumni of St John's College, Cambridge]]
[[Category:Members of the European Molecular Biology Organization]]
[[Category:Fellows of King's College, Cambridge]]
[[Category:Academics of the University of Cambridge]]
[[Category:Commanders of the Order of the British Empire]]
[[Category:English biochemists]]
[[Category:British agnostics]]
[[Category:Fellows of the Royal Society]]
[[Category:Foreign associates of the National Academy of Sciences]]
[[Category:Members of the Order of Merit]]
[[Category:Members of the Order of the Companions of Honour]]
[[Category:Nobel laureates in Chemistry]]
[[Category:Nobel laureates with multiple Nobel awards]]
[[Category:British Nobel laureates]]
[[Category:People educated at Bryanston School]]
[[Category:People educated at Malvern College]]
[[Category:Members of the French Academy of Sciences]]
[[Category:English pacifists]]
[[Category:British conscientious objectors]]
[[Category:People educated at The Downs School, Herefordshire]]
[[Category:Royal Medal winners]]
[[Category:Recipients of the Copley Medal]]
[[Category:English Nobel laureates]]
[[Category:People from Cotswold District]]
[[Category:Recipients of the Albert Lasker Award for Basic Medical Research]]
[[Category:Fellows of the Australian Academy of Science]]
[[Category:Foreign fellows of the Indian National Science Academy]]
[[Category:World Constitutional Convention call signatories]]