Frederick Sanger

Frederick Sanger OM CH CBE FRS FAA (/ˈsæŋər/; 13

August 1918 – 19 November 2013) was a British Frederick Sanger
OM CH CBE FRS FAA
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 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 Born 13 August 1918
with Walter Gilbert and Paul Berg. Rendcomb, Gloucestershire,
England
He is one of only three people to have won multiple
Nobel Prizes in the same category (the others being Died 19 November 2013 (aged 95)
John Bardeen in physics and Karl Barry Sharpless in Cambridge, England[3]
chemistry),[5] and one of five persons with two Nobel Alma mater St John's College, Cambridge
Prizes. (BA, PhD)
Known for Determining the amino acid
sequence of insulin
Early life and education Sanger sequencing
Sanger Centre
Frederick Sanger was born on 13 August 1918 in
Rendcomb, a small village in Gloucestershire, Spouse Margaret Joan Howe[4]
England, the second son of Frederick Sanger, a general Awards Nobel Prize in Chemistry (1958,
practitioner, and his wife, Cicely Sanger (née 1980)
Crewdson).[6] He was one of three children. His
Foreign Associate of the National
brother, Theodore, was only a year older, while his
Academy of Sciences (1967)
sister May (Mary) was five years younger.[7] His father
Royal Medal (1969)
had worked as an Anglican medical missionary in
China but returned to England because of ill health. He Gairdner Foundation
was 40 in 1916 when he married Cicely, who was four International Award (1971)
years younger. Sanger's father converted to Quakerism William Bate Hardy Prize (1976)
Copley Medal (1977)
soon after his two sons were born and brought up the Louisa Gross Horwitz Prize
children as Quakers. Sanger's mother was the daughter (1979)
of an affluent cotton manufacturer and had a Quaker Scientific career
background, but was not a Quaker.[7]
Fields Biochemistry
When Sanger was around five years old the family Institutions University of Cambridge
moved to the small village of Tanworth-in-Arden in Laboratory of Molecular Biology
Warwickshire. The family was reasonably wealthy and
employed a governess to teach the children. In 1927, at Thesis The metabolism of the amino
acid lysine in the animal body (htt
the age of nine, he was sent to the Downs School, a
p://idiscover.lib.cam.ac.uk/primo-
residential preparatory school run by Quakers near
explore/fulldisplay?docid=44CAM
Malvern. His brother Theo was a year ahead of him at
_ALMA21428208290003606&co
the same school. In 1932, at the age of 14, he was sent
ntext=L&vid=44CAM_PROD&se
to the recently established Bryanston School in Dorset.
arch_scope=default_scope&tab=
This used the Dalton system and had a more liberal
default_tab&lang=en_US) (1943)
regime which Sanger much preferred. At the school he
liked his teachers and particularly enjoyed scientific Doctoral Albert Neuberger[1]
subjects.[7] Able to complete his School Certificate a advisor
year early, for which he was awarded seven credits, Doctoral George Brownlee[2]
Sanger was able to spend most of his last year of students Elizabeth Blackburn
school experimenting in the laboratory alongside his
Rodney Porter
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.[8] 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.[9]

In 1936 Sanger went to St John's College, Cambridge, to study natural sciences. His father had attended
the same college. For Part 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 I. For his Part 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, Joseph Needham and Ernest Baldwin.[7]

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 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 has nothing but respect for the religious and states he took two
things from it, truth and respect for all life.[10] 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.[7]

Sanger began studying for a PhD in October 1940 under 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.[7] Sanger changed his research project to study the
metabolism of lysine[11] and a more practical problem concerning the nitrogen of potatoes.[12] His thesis
had the title, "The metabolism of the amino acid lysine in the animal body". He was examined by Charles
Harington and Albert Charles Chibnall and awarded his doctorate in 1943.[7]

Research and career

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.[13] Chibnall had already done some work on the amino
acid composition of bovine insulin[14] and suggested that Sanger look at the amino groups in the protein.
Insulin could be purchased from the pharmacy chain 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 and then from 1944 until 1951 by a Beit
Memorial Fellowship for Medical Research.[6]

Sanger's first triumph was to determine the complete amino acid sequence of the two polypeptide chains
of bovine insulin, A and B, in 1952 and 1951, respectively.[15][16] Prior to this it was widely assumed that
proteins were somewhat amorphous. In determining these sequences, Sanger proved that proteins have a
defined chemical composition.[7]

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.[17] 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
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.[7]
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 sequences to deduce the complete
structure of insulin. Finally, because the A and B chains are
physiologically inactive without the three linking disulfide bonds
(two interchain, one intrachain on A), Sanger and coworkers
determined their assignments in 1955.[18][19] 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.[20] This
discovery was crucial to the later sequence hypothesis of Francis
Crick for developing ideas of how DNA codes for proteins.[21]

Sequencing RNA
From 1951 Sanger was a member of the external staff of the
Medical Research Council[6] 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.[7]

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.[21]
Amino acid sequence of bovine
insulin, with disulfide bridges shown
The key challenge in the work was finding a pure piece of RNA to
in red.
sequence. In the course of the work he discovered in 1964, with
Kjeld Marcker, the formylmethionine tRNA which initiates protein
synthesis in bacteria.[22] He was beaten in the race to be the first to sequence a tRNA molecule by a group
led by Robert Holley from Cornell University, who published the sequence of the 77 ribonucleotides of
alanine tRNA from Saccharomyces cerevisiae in 1965.[23] By 1967 Sanger's group had determined the
nucleotide sequence of the 5S ribosomal RNA from Escherichia coli, a small RNA of 120 nucleotides.[24]

Sequencing DNA
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.[25] 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.[26][27] 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 φX174.[28] This was the first fully sequenced DNA-based genome. To their
surprise they discovered that the coding regions of some of the genes overlapped with one another.[2]

In 1977 Sanger and colleagues introduced the "dideoxy" chain-termination method for sequencing DNA
molecules, also known as the "Sanger method".[27][29] 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.[30] The new method was used by
Sanger and colleagues to sequence human mitochondrial DNA (16,569 base pairs)[31] and bacteriophage
λ (48,502 base pairs).[32] The dideoxy method was eventually used to sequence the entire human
genome.[33]

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 Porter who joined the research group in
1947.[2] Porter later shared the 1972 Nobel Prize in Physiology or Medicine with Gerald Edelman for his
work on the chemical structure of antibodies.[34] Elizabeth Blackburn studied for a PhD in Sanger's
laboratory between 1971 and 1974.[2][35] She shared the 2009 Nobel Prize in Physiology or Medicine
with Carol W. Greider and Jack W. Szostak for her work on telomeres and the action of telomerase.[36]

Sanger's rule

... anytime you get technical development that’s two to threefold or more efficient, accurate,
cheaper, a whole range of experiments opens up.[37]

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 (Physics, 1903 and Chemistry, 1911), Linus Pauling
(Chemistry, 1954 and Peace, 1962), John Bardeen (twice Physics, 1956 and 1972), and Karl Barry
Sharpless (twice Chemistry, 2001 and 2022).[5]

Elected Fellow of the Royal Society (FRS) in 1954[2]

Commander of the Order of the British Empire (CBE) – 1963[2]
Member of the Order of the Companions of Honour (CH) – 1981[2]
Member of the Order of Merit (OM) – 1986[2]
Corresponding Member of the Australian Academy of Science – 1982[2]
William Bate Hardy Prize – 1976[2]
Nobel Prize in Chemistry – 1958, 1980[20][30]
Corday–Morgan Medal – 1951[2]
Royal Medal – 1969[2]
 Gairdner Foundation International Award – 1971[2]
Copley Medal – 1977[2]
G.W. Wheland Award – 1978[2]
Louisa Gross Horwitz Prize of Columbia University – 1979[2]
Albert Lasker Award for Basic Medical Research – 1979[2]
Association of Biomolecular Resource Facilities Award – 1994[38]
Golden Plate Award of the American Academy of Achievement – 2000[39][40]
Citation for Chemical Breakthrough Award from the Division of History of Chemistry of the
American Chemical Society – 2016[41][42][29]
The Wellcome Trust Sanger Institute (formerly the Sanger Centre) is named in his honour.[2]

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.[6] He said that his wife had "contributed more to his work than anyone
else by providing a peaceful and happy home."[43]

Later life
Sanger retired in 1983, aged 65, to his home, "Far
Leys", in Swaffham Bulbeck outside Cambridge.[2]

In 1992, the Wellcome Trust and the Medical

Research Council founded the Sanger Centre (now
The Sanger Institute
the Sanger Institute), named after him.[44] 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."[44] 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 sequencing of the human genome.[44] 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.[45] In an interview published in
the 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."[46]

He declined the offer of a 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, which can have only 24 living members.[43][45][46]
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 noted that Sanger, "the most self-effacing person you could hope to meet",
was spending his time gardening at his Cambridgeshire home.[47]

Sanger died in his sleep at Addenbrooke's Hospital in Cambridge on 19 November 2013.[43][48] As noted
in his obituary, he had described himself as "just a chap who messed about in a lab",[49] and
"academically not brilliant".[50]

Global policy
He was one of the signatories of the agreement to convene a convention for drafting a world
constitution.[51][52] 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.[53]

Selected publications
Neuberger, A.; Sanger, F. (1942), "The nitrogen of the potato", Biochemical Journal, 36 (7–
9): 662–671, doi:10.1042/bj0360662 (https://doi.org/10.1042%2Fbj0360662), PMC 1266851
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1266851), PMID 16747571 (https://pubmed.
ncbi.nlm.nih.gov/16747571).
Neuberger, A.; Sanger, F. (1944), "The metabolism of lysine", Biochemical Journal, 38 (1):
119–125, doi:10.1042/bj0380119 (https://doi.org/10.1042%2Fbj0380119), PMC 1258037 (ht
tps://www.ncbi.nlm.nih.gov/pmc/articles/PMC1258037), PMID 16747737 (https://pubmed.nc
bi.nlm.nih.gov/16747737).
Sanger, F. (1945), "The free amino groups of insulin", Biochemical Journal, 39 (5): 507–515,
doi:10.1042/bj0390507 (https://doi.org/10.1042%2Fbj0390507), PMC 1258275 (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC1258275), PMID 16747948 (https://pubmed.ncbi.nlm.nih.
gov/16747948).
Sanger, F. (1947), "Oxidation of insulin by performic acid", Nature, 160 (4061): 295–296,
Bibcode:1947Natur.160..295S (https://ui.adsabs.harvard.edu/abs/1947Natur.160..295S),
doi:10.1038/160295b0 (https://doi.org/10.1038%2F160295b0), PMID 20344639 (https://pub
med.ncbi.nlm.nih.gov/20344639), S2CID 4127677 (https://api.semanticscholar.org/CorpusI
D:4127677).
Porter, R.R.; Sanger, F. (1948), "The free amino groups of haemoglobins", Biochemical
Journal, 42 (2): 287–294, doi:10.1042/bj0420287 (https://doi.org/10.1042%2Fbj0420287),
PMC 1258669 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1258669), PMID 16748281
(https://pubmed.ncbi.nlm.nih.gov/16748281).
Sanger, F. (1949a), "Fractionation of oxidized insulin", Biochemical Journal, 44 (1): 126–
128, doi:10.1042/bj0440126 (https://doi.org/10.1042%2Fbj0440126), PMC 1274818 (https://
www.ncbi.nlm.nih.gov/pmc/articles/PMC1274818), PMID 16748471 (https://pubmed.ncbi.nl
m.nih.gov/16748471).
Sanger, F. (1949b), "The terminal peptides of insulin", Biochemical Journal, 45 (5): 563–574,
doi:10.1042/bj0450563 (https://doi.org/10.1042%2Fbj0450563), PMC 1275055 (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC1275055), PMID 15396627 (https://pubmed.ncbi.nlm.nih.
gov/15396627).
Sanger, F.; Tuppy, H. (1951a), "The amino-acid sequence in the phenylalanyl chain of
insulin. 1. The identification of lower peptides from partial hydrolysates", Biochemical
Journal, 49 (4): 463–481, doi:10.1042/bj0490463 (https://doi.org/10.1042%2Fbj0490463),
PMC 1197535 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1197535), PMID 14886310
(https://pubmed.ncbi.nlm.nih.gov/14886310).
Sanger, F.; Tuppy, H. (1951b), "The amino-acid sequence in the phenylalanyl chain of
insulin. 2. The investigation of peptides from enzymic hydrolysates", Biochemical Journal,
49 (4): 481–490, doi:10.1042/bj0490481 (https://doi.org/10.1042%2Fbj0490481),
PMC 1197536 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1197536), PMID 14886311
(https://pubmed.ncbi.nlm.nih.gov/14886311).
Sanger, F.; Thompson, E.O.P. (1953a), "The amino-acid sequence in the glycyl chain of
insulin. 1. The identification of lower peptides from partial hydrolysates", Biochemical
Journal, 53 (3): 353–366, doi:10.1042/bj0530353 (https://doi.org/10.1042%2Fbj0530353),
PMC 1198157 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1198157), PMID 13032078
(https://pubmed.ncbi.nlm.nih.gov/13032078).
Sanger, F.; Thompson, E.O.P. (1953b), "The amino-acid sequence in the glycyl chain of
insulin. 2. The investigation of peptides from enzymic hydrolysates", Biochemical Journal,
53 (3): 366–374, doi:10.1042/bj0530366 (https://doi.org/10.1042%2Fbj0530366),
PMC 1198158 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1198158), PMID 13032079
(https://pubmed.ncbi.nlm.nih.gov/13032079).
Sanger, F.; Thompson, E.O.P.; Kitai, R. (1955), "The amide groups of insulin", Biochemical
Journal, 59 (3): 509–518, doi:10.1042/bj0590509 (https://doi.org/10.1042%2Fbj0590509),
PMC 1216278 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1216278), PMID 14363129
(https://pubmed.ncbi.nlm.nih.gov/14363129).
Ryle, A.P.; Sanger, F.; Smith, L.F.; Kitai, R. (1955), "The disulphide bonds of insulin",
Biochemical Journal, 60 (4): 541–556, doi:10.1042/bj0600541 (https://doi.org/10.1042%2Fbj
0600541), PMC 1216151 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1216151),
PMID 13249947 (https://pubmed.ncbi.nlm.nih.gov/13249947).
Brown, H.; Sanger, F.; Kitai, R. (1955), "The structure of pig and sheep insulins",
Biochemical Journal, 60 (4): 556–565, doi:10.1042/bj0600556 (https://doi.org/10.1042%2Fbj
0600556), PMC 1216152 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1216152),
PMID 13249948 (https://pubmed.ncbi.nlm.nih.gov/13249948).
Sanger, F. (1959), "Chemistry of Insulin: determination of the structure of insulin opens the
way to greater understanding of life processes", Science, 129 (3359): 1340–1344,
Bibcode:1959Sci...129.1340G (https://ui.adsabs.harvard.edu/abs/1959Sci...129.1340G),
doi:10.1126/science.129.3359.1340 (https://doi.org/10.1126%2Fscience.129.3359.1340),
PMID 13658959 (https://pubmed.ncbi.nlm.nih.gov/13658959).
Milstein, C.; Sanger, F. (1961), "An amino acid sequence in the active centre of
phosphoglucomutase", Biochemical Journal, 79 (3): 456–469, doi:10.1042/bj0790456 (http
s://doi.org/10.1042%2Fbj0790456), PMC 1205670 (https://www.ncbi.nlm.nih.gov/pmc/article
s/PMC1205670), PMID 13771000 (https://pubmed.ncbi.nlm.nih.gov/13771000).
Marcker, K.; Sanger, F. (1964), "N-formyl-methionyl-S-RNA", Journal of Molecular Biology, 8
(6): 835–840, doi:10.1016/S0022-2836(64)80164-9 (https://doi.org/10.1016%2FS0022-283
6%2864%2980164-9), PMID 14187409 (https://pubmed.ncbi.nlm.nih.gov/14187409).
Sanger, F.; Brownlee, G.G.; Barrell, B.G. (1965), "A two-dimensional fractionation procedure
for radioactive nucleotides", Journal of Molecular Biology, 13 (2): 373–398,
doi:10.1016/S0022-2836(65)80104-8 (https://doi.org/10.1016%2FS0022-2836%2865%2980
104-8), PMID 5325727 (https://pubmed.ncbi.nlm.nih.gov/5325727).
Brownlee, G.G.; Sanger, F.; Barrell, B.G. (1967), "Nucleotide sequence of 5S-ribosomal
RNA from Escherichia coli", Nature, 215 (5102): 735–736, Bibcode:1967Natur.215..735B (ht
tps://ui.adsabs.harvard.edu/abs/1967Natur.215..735B), doi:10.1038/215735a0 (https://doi.or
g/10.1038%2F215735a0), PMID 4862513 (https://pubmed.ncbi.nlm.nih.gov/4862513),
S2CID 4270186 (https://api.semanticscholar.org/CorpusID:4270186).
Brownlee, G.G.; Sanger, F. (1967), "Nucleotide sequences from the low molecular weight
ribosomal RNA of Escherichia coli", Journal of Molecular Biology, 23 (3): 337–353,
doi:10.1016/S0022-2836(67)80109-8 (https://doi.org/10.1016%2FS0022-2836%2867%2980
109-8), PMID 4291728 (https://pubmed.ncbi.nlm.nih.gov/4291728).
Brownlee, G.G.; Sanger, F.; Barrell, B.G. (1968), "The sequence of 5S ribosomal ribonucleic
acid", Journal of Molecular Biology, 34 (3): 379–412, doi:10.1016/0022-2836(68)90168-X (ht
tps://doi.org/10.1016%2F0022-2836%2868%2990168-X), PMID 4938553 (https://pubmed.n
cbi.nlm.nih.gov/4938553).
Adams, J.M.; Jeppesen, P.G.; Sanger, F.; Barrell, B.G. (1969), "Nucleotide sequence from
the coat protein cistron of R17 bacteriophage RNA", Nature, 223 (5210): 1009–1014,
Bibcode:1969Natur.223.1009A (https://ui.adsabs.harvard.edu/abs/1969Natur.223.1009A),
doi:10.1038/2231009a0 (https://doi.org/10.1038%2F2231009a0), PMID 5811898 (https://pu
bmed.ncbi.nlm.nih.gov/5811898), S2CID 4152602 (https://api.semanticscholar.org/CorpusI
D:4152602).
Barrell, B.G.; Sanger, F. (1969), "The sequence of phenylalanine tRNA from E. coli", FEBS
Letters, 3 (4): 275–278, Bibcode:1969FEBSL...3..275B (https://ui.adsabs.harvard.edu/abs/1
969FEBSL...3..275B), doi:10.1016/0014-5793(69)80157-2 (https://doi.org/10.1016%2F0014
-5793%2869%2980157-2), PMID 11947028 (https://pubmed.ncbi.nlm.nih.gov/11947028),
S2CID 34155866 (https://api.semanticscholar.org/CorpusID:34155866).
Jeppesen, P.G.; Barrell, B.G.; Sanger, F.; Coulson, A.R. (1972), "Nucleotide sequences of
two fragments from the coat-protein cistron of bacteriophage R17 ribonucleic acid",
Biochemical Journal, 128 (5): 993–1006, doi:10.1042/bj1280993h (https://doi.org/10.1042%
2Fbj1280993h), PMC 1173988 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1173988),
PMID 4566195 (https://pubmed.ncbi.nlm.nih.gov/4566195).
Sanger, F.; Donelson, J.E.; Coulson, A.R.; Kössel, H.; Fischer, D. (1973), "Use of DNA
Polymerase I Primed by a Synthetic Oligonucleotide to Determine a Nucleotide Sequence in
Phage f1 DNA", Proceedings of the National Academy of Sciences USA, 70 (4): 1209–1213,
Bibcode:1973PNAS...70.1209S (https://ui.adsabs.harvard.edu/abs/1973PNAS...70.1209S),
doi:10.1073/pnas.70.4.1209 (https://doi.org/10.1073%2Fpnas.70.4.1209), PMC 433459 (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC433459), PMID 4577794 (https://pubmed.ncbi.n
lm.nih.gov/4577794).
Sanger, F.; Coulson, A.R. (1975), "A rapid method for determining sequences in DNA by
primed synthesis with DNA polymerase", Journal of Molecular Biology, 94 (3): 441–448,
doi:10.1016/0022-2836(75)90213-2 (https://doi.org/10.1016%2F0022-2836%2875%299021
3-2), PMID 1100841 (https://pubmed.ncbi.nlm.nih.gov/1100841).
Sanger, F.; Nicklen, S.; Coulson, A.R. (1977), "DNA sequencing with chain-terminating
inhibitors", Proceedings of the National Academy of Sciences USA, 74 (12): 5463–5467,
Bibcode:1977PNAS...74.5463S (https://ui.adsabs.harvard.edu/abs/1977PNAS...74.5463S),
doi:10.1073/pnas.74.12.5463 (https://doi.org/10.1073%2Fpnas.74.12.5463), PMC 431765
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC431765), PMID 271968 (https://pubmed.ncb
i.nlm.nih.gov/271968). According to the Institute for Scientific Information (ISI) database, by
October 2010 this paper had been cited over 64,000 times.
Sanger, F.; Air, G.M.; Barrell, B.G.; Brown, N.L.; Coulson, A.R.; Fiddes, C.A.; Hutchinson,
C.A.; Slocombe, P.M.; Smith, M. (1977), "Nucleotide sequence of bacteriophage φX174
DNA", Nature, 265 (5596): 687–695, Bibcode:1977Natur.265..687S (https://ui.adsabs.harvar
d.edu/abs/1977Natur.265..687S), doi:10.1038/265687a0 (https://doi.org/10.1038%2F26568
7a0), PMID 870828 (https://pubmed.ncbi.nlm.nih.gov/870828), S2CID 4206886 (https://api.s
emanticscholar.org/CorpusID:4206886).
Sanger, F.; Coulson, A.R. (1978), "The use of thin acrylamide gels for DNA sequencing",
FEBS Letters, 87 (1): 107–110, Bibcode:1978FEBSL..87..107S (https://ui.adsabs.harvard.e
du/abs/1978FEBSL..87..107S), doi:10.1016/0014-5793(78)80145-8 (https://doi.org/10.101
6%2F0014-5793%2878%2980145-8), PMID 631324 (https://pubmed.ncbi.nlm.nih.gov/6313
24), S2CID 1620755 (https://api.semanticscholar.org/CorpusID:1620755).
 Sanger, F.; Coulson, A.R.; Barrell, B.G.; Smith, A.J.; Roe, B.A. (1980), "Cloning in single-
stranded bacteriophage as an aid to rapid DNA sequencing", Journal of Molecular Biology,
143 (2): 161–178, doi:10.1016/0022-2836(80)90196-5 (https://doi.org/10.1016%2F0022-283
6%2880%2990196-5), PMID 6260957 (https://pubmed.ncbi.nlm.nih.gov/6260957).
Anderson, S.; Bankier, A.T.; Barrell, B.G.; De Bruijn, M.H.; Coulson, A.R.; Drouin, J.; Eperon,
I.C.; Nierlich, D.P.; Roe, B.A.; Sanger, F.; Schreier, P.H.; Smith, A.J.; Staden, R.; Young, I.G.
(1981), "Sequence and organization of the human mitochondrial genome", Nature, 290
(5806): 457–465, Bibcode:1981Natur.290..457A (https://ui.adsabs.harvard.edu/abs/1981Nat
ur.290..457A), doi:10.1038/290457a0 (https://doi.org/10.1038%2F290457a0),
PMID 7219534 (https://pubmed.ncbi.nlm.nih.gov/7219534), S2CID 4355527 (https://api.sem
anticscholar.org/CorpusID:4355527).
Anderson, S.; De Bruijn, M.H.; Coulson, A.R.; Eperon, I.C.; Sanger, F.; Young, I.G. (1982),
"Complete sequence of bovine mitochondrial DNA. Conserved features of the mammalian
mitochondrial genome", Journal of Molecular Biology, 156 (4): 683–717, doi:10.1016/0022-
2836(82)90137-1 (https://doi.org/10.1016%2F0022-2836%2882%2990137-1),
PMID 7120390 (https://pubmed.ncbi.nlm.nih.gov/7120390).
Sanger, F.; Coulson, A.R.; Hong, G.F.; Hill, D.F.; Petersen, G.B. (1982), "Nucleotide
sequence of bacteriophage λ DNA", Journal of Molecular Biology, 162 (4): 729–773,
doi:10.1016/0022-2836(82)90546-0 (https://doi.org/10.1016%2F0022-2836%2882%299054
6-0), PMID 6221115 (https://pubmed.ncbi.nlm.nih.gov/6221115).
Sanger, F. (1988), "Sequences, sequences, and sequences", Annual Review of
Biochemistry, 57: 1–28, doi:10.1146/annurev.bi.57.070188.000245 (https://doi.org/10.114
6%2Fannurev.bi.57.070188.000245), PMID 2460023 (https://pubmed.ncbi.nlm.nih.gov/2460
023).

References

Citations
1. Allen, A.K.; Muir, H.M. (2001). "Albert Neuberger. 15 April 1908 – 14 August 1996".
Biographical Memoirs of Fellows of the Royal Society. 47: 369–382.
doi:10.1098/rsbm.2001.0021 (https://doi.org/10.1098%2Frsbm.2001.0021). JSTOR 770373
(https://www.jstor.org/stable/770373). PMID 15124648 (https://pubmed.ncbi.nlm.nih.gov/151
24648). S2CID 72943723 (https://api.semanticscholar.org/CorpusID:72943723).
2. Brownlee, George G. (2015). "Frederick Sanger CBE CH OM. 13 August 1918 – 19
November 2013" (https://doi.org/10.1098%2Frsbm.2015.0013). Biographical Memoirs of
Fellows of the Royal Society. 61: 437–466. doi:10.1098/rsbm.2015.0013 (https://doi.org/10.
1098%2Frsbm.2015.0013).
3. "Seven days: 22–28 November 2013" (https://doi.org/10.1038%2F503442a). Nature. 503
(7477): 442–443. 2013. Bibcode:2013Natur.503..442. (https://ui.adsabs.harvard.edu/abs/20
13Natur.503..442.). doi:10.1038/503442a (https://doi.org/10.1038%2F503442a).
4. "The Nobel Prize in Chemistry 1958" (https://www.nobelprize.org/prizes/chemistry/1958/san
ger/biographical/).
5. "Nobel Prize Facts" (https://www.nobelprize.org/nobel_prizes/facts/). Nobelprize.org.
Retrieved 1 September 2015.
6. "The Nobel Prize in Chemistry 1958: Frederick Sanger – biography" (https://www.nobelpriz
e.org/prizes/chemistry/1958/sanger/biographical/). Nobelprize.org. Retrieved 10 August
2020.
 7. "A Life of Research on the Sequences of Proteins and Nucleic Acids: Fred Sanger in
conversation with George Brownlee" (https://web.archive.org/web/20140313164607/http://jis
cmediahub.ac.uk/record/display/012-00002584). Biochemical Society, Edina – Film & Sound
Online. 9 October 1992. Archived from the original (http://jiscmediahub.ac.uk/record/display/
012-00002584) on 13 March 2014. Retrieved 29 April 2013.. Subscription required. A
200 min interview divided into 44 segments. Notes give the content of each segment.
8. Marks, Lara. "Sanger's early life: From the cradle to the laboratory" (http://www.whatisbiotec
hnology.org/exhibitions/sanger/early). The path to DNA sequencing: The life and work of
Fred Sanger. What is Biotechnology. Retrieved 1 September 2015.
9. Jeffers, Joe S. (2017). Frederick Sanger Two-Time Nobel Laureate in Chemistry. Springer
International. ISBN 978-3-319-54707-7.
10. "The Nobel Prize in Chemistry 1980" (https://www.nobelprize.org/prizes/chemistry/1980/san
ger/25898-interview-transcript-1980-2/).
11. Sanger, Frederick (1944). The metabolism of the amino acid lysine in the animal body (http
s://web.archive.org/web/20210417201357/http://ulmss-newton.lib.cam.ac.uk/vwebv/holdings
Info?bibId=35253) (PhD thesis). University of Cambridge. Archived from the original (http://ul
mss-newton.lib.cam.ac.uk/vwebv/holdingsInfo?bibId=35253) on 17 April 2021. Retrieved
5 December 2013.
12. Neuberger & Sanger 1942; Neuberger & Sanger 1944
13. "Frederick Sanger, Ph.D. Biography and Interview" (https://achievement.org/achiever/frederi
ck-sanger-ph-d/#interview). www.achievement.org. American Academy of Achievement.
14. Chibnall, A. C. (1942). "Bakerian Lecture: Amino-Acid Analysis and the Structure of
Proteins" (http://rspa.royalsocietypublishing.org/content/royprsa/181/985/210.full.pdf) (PDF).
Proceedings of the Royal Society B: Biological Sciences. 131 (863): 136–160.
Bibcode:1942RSPSB.131..136C (https://ui.adsabs.harvard.edu/abs/1942RSPSB.131..136
C). doi:10.1098/rspb.1942.0021 (https://doi.org/10.1098%2Frspb.1942.0021).
S2CID 85124201 (https://api.semanticscholar.org/CorpusID:85124201). Section on insulin
starts on page 153.
15. Sanger & Tuppy 1951a; Sanger & Tuppy 1951b; Sanger & Thompson 1953a; Sanger &
Thompson 1953b
16. Sanger, F. (1958), Nobel lecture: The chemistry of insulin (http://nobelprize.org/nobel_prize
s/chemistry/laureates/1958/sanger-lecture.pdf) (PDF), Nobelprize.org, retrieved 18 October
2010. Sanger's Nobel lecture was also published in Science: Sanger 1959
17. Marks, Lara. "Sequencing proteins: Insulin" (http://www.whatisbiotechnology.org/exhibitions/
sanger/insulin). The path to DNA sequencing: The life and work of Fred Sanger. What is
Biotechnology. Retrieved 1 September 2015.
18. Ryle et al. 1955.
19. Stretton, A.O. (2002). "The first sequence. Fred Sanger and insulin" (https://www.ncbi.nlm.ni
h.gov/pmc/articles/PMC1462286). Genetics. 162 (2): 527–532.
doi:10.1093/genetics/162.2.527 (https://doi.org/10.1093%2Fgenetics%2F162.2.527).
PMC 1462286 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1462286). PMID 12399368
(https://pubmed.ncbi.nlm.nih.gov/12399368).
20. "The Nobel Prize in Chemistry 1958: Frederick Sanger" (http://nobelprize.org/nobel_prizes/c
hemistry/laureates/1958/). Nobelprize.org. Retrieved 8 October 2010.
21. Marks, Lara. "The path to sequencing nucleic acids" (http://www.whatisbiotechnology.org/ex
hibitions/sanger/path). The path to DNA sequencing: The life and work of Fred Sanger.
What is Biotechnology. Retrieved 1 September 2015.
22. Marcker & Sanger 1964
23. Holley, R. W.; Apgar, J.; Everett, G. A.; Madison, J. T.; Marquisee, M.; Merrill, S. H.;
Penswick, J. R.; Zamir, A. (1965). "Structure of a Ribonucleic Acid". Science. 147 (3664):
1462–1465. Bibcode:1965Sci...147.1462H (https://ui.adsabs.harvard.edu/abs/1965Sci...14
7.1462H). doi:10.1126/science.147.3664.1462 (https://doi.org/10.1126%2Fscience.147.366
4.1462). PMID 14263761 (https://pubmed.ncbi.nlm.nih.gov/14263761). S2CID 40989800 (ht
tps://api.semanticscholar.org/CorpusID:40989800).
24. Brownlee, Sanger & Barrell 1967; Brownlee, Sanger & Barrell 1968
25. Sanger et al. 1973
26. Sanger & Coulson 1975
27. Sanger, F. (1980). "Nobel lecture: Determination of nucleotide sequences in DNA" (https://w
ww.nobelprize.org/uploads/2018/06/sanger-lecture-1.pdf) (PDF). Nobelprize.org. Retrieved
15 September 2019.
28. Sanger et al. 1977
29. Sanger, Nicklen & Coulson 1977.
30. "The Nobel Prize in Chemistry 1980: Paul Berg, Walter Gilbert, Frederick Sanger" (http://nob
elprize.org/nobel_prizes/chemistry/laureates/1980/). Nobelprize.org. Retrieved 8 October
2010.
31. Anderson et al. 1981
32. Sanger et al. 1982
33. Walker, John (2014). "Frederick Sanger (1918–2013) Double Nobel-prizewinning genomics
pioneer" (https://doi.org/10.1038%2F505027a). Nature. 505 (7481): 27.
Bibcode:2014Natur.505...27W (https://ui.adsabs.harvard.edu/abs/2014Natur.505...27W).
doi:10.1038/505027a (https://doi.org/10.1038%2F505027a). PMID 24380948 (https://pubme
d.ncbi.nlm.nih.gov/24380948).
34. "The Nobel Prize in Physiology or Medicine 1972" (https://www.nobelprize.org/nobel_prizes/
medicine/laureates/1972/). Nobelprize.org. Retrieved 1 September 2015.
35. Blackburn, E. H. (1974). Sequence studies on bacteriophage ØX174 DNA by transcription
(https://web.archive.org/web/20160305133053/http://ethos.bl.uk/OrderDetails.do?uin=uk.bl.
ethos.449954) (PhD thesis). University of Cambridge. Archived from the original (http://etho
s.bl.uk/OrderDetails.do?uin=uk.bl.ethos.449954) on 5 March 2016. Retrieved 4 July 2015.
36. "The Nobel Prize in Physiology or Medicine 2009" (https://www.nobelprize.org/nobel_prizes/
medicine/laureates/2009/). Nobelprize.org. Retrieved 1 September 2015.
37. "Schlessinger, David" (https://www.genome.gov/sites/default/files/media/files/2019-05/david
_schlessinger_transcript.pdf) (PDF). National Human Genome Research Institute (NHGRI,
genome.gov). March 2018.
38. "The ABRF Award for Outstanding Contributions to Biomolecular Technologies" (https://web.
archive.org/web/20200806131051/https://abrf.org/awards/abrf-award-outstanding-contributi
ons-biomolecular-technologies). Association of Biomolecular Resource facilities. Archived
from the original (https://abrf.org/awards/abrf-award-outstanding-contributions-biomolecular-
technologies) on 6 August 2020. Retrieved 11 August 2020.
39. "Golden Plate Awardees of the American Academy of Achievement" (https://www.achieveme
nt.org/our-history/golden-plate-awards/#science-exploration/). www.achievement.org.
American Academy of Achievement.
40. "Summit Overview Photo" (https://achievement.org/summit/). "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."
41. "2016 Awardees" (https://web.archive.org/web/20170506160732/http://www.scs.illinois.edu/
~mainzv/HIST/awards/CCB-2016_Awardees.php). American Chemical Society, Division of
the History of Chemistry. University of Illinois at Urbana-Champaign School of Chemical
Sciences. 2016. Archived from the original (http://www.scs.illinois.edu/~mainzv/HIST/award
s/CCB-2016_Awardees.php) on 6 May 2017. Retrieved 14 June 2017.
42. "Citation for Chemical Breakthrough Award" (http://www.scs.illinois.edu/~mainzv/HIST/awar
ds/Citations/2016-ACS%20Rendering_Sanger.pdf) (PDF). American Chemical Society,
Division of the History of Chemistry. University of Illinois at Urbana-Champaign School of
Chemical Sciences. 2016. Retrieved 14 June 2017.
43. "Frederick Sanger, OM" (https://www.telegraph.co.uk/news/obituaries/science-obituaries/10
462574/Frederick-Sanger-OM.html). The Telegraph. 20 November 2013. Archived (https://g
hostarchive.org/archive/20220112/https://www.telegraph.co.uk/news/obituaries/science-obit
uaries/10462574/Frederick-Sanger-OM.html) from the original on 12 January 2022.
Retrieved 20 November 2013.
44. "Frederick Sanger" (https://web.archive.org/web/20110407064631/http://www.sanger.ac.uk/
about/people/biographies/fsanger.html). Wellcome Trust Sanger Institute. Archived from the
original (http://www.sanger.ac.uk/about/people/biographies/fsanger.html) on 7 April 2011.
Retrieved 12 October 2010.
45. Hargittai, István (April 1999). "Interview: Frederick Sanger". The Chemical Intelligencer. 4
(2). New York: Springer-Verlag: 6–11.. This interview, which took place on 16 September
1997, was republished in Hargittai, István (2002). "Chapter 5: Frederick Sanger". Candid
science II: conversations with famous biomedical scientists. London: Imperial College Press.
pp. 73–83. ISBN 978-1-86094-288-4.
46. Ahuja, Anjana (12 January 2000). "The double Nobel laureate who began the book of life" (h
ttps://web.archive.org/web/20081211053639/http://www.warwick.ac.uk/fac/sci/Chemistry/Me
dChem/MedChemInfo/info/Sanger.html). The Times. London. p. 40. Archived from the
original (http://www.warwick.ac.uk/fac/sci/Chemistry/MedChem/MedChemInfo/info/Sanger.ht
ml) on 11 December 2008. Retrieved 18 October 2010 – via warwick.ac.uk.
47. Bhattachjee, Yudhijit, ed. (2007). "Newsmakers: A Life in Science" (http://www.sciencemag.o
rg/content/vol317/issue5840/newsmakers.dtl?cookietest=yes). Science. 317 (5840): 879.
doi:10.1126/science.317.5840.879e (https://doi.org/10.1126%2Fscience.317.5840.879e).
S2CID 220092058 (https://api.semanticscholar.org/CorpusID:220092058).
48. "Frederick Sanger: Nobel Prize winner dies at 95" (https://www.bbc.co.uk/news/science-envi
ronment-25020112). BBC.co.uk. 20 November 2013. Retrieved 20 November 2013.
49. "Frederick Sanger: Unassuming British biochemist whose pivotal and far-reaching
discoveries made him one of a handful of double Nobel prizewinners". The Times. London.
21 November 2013. p. 63.
50. "Frederick Sanger's achievements cannot be overstated" (https://theconversation.com/frede
rick-sangers-achievements-cannot-be-overstated-20596). The Conversation. 21 November
2013.
51. "Letters from Thane Read asking Helen Keller to sign the World Constitution for world
peace. 1961" (https://www.afb.org/HelenKellerArchive?a=d&d=A-HK01-07-B149-F04-022.1.
8). Helen Keller Archive. American Foundation for the Blind. Retrieved 1 July 2023.
52. "Letter from World Constitution Coordinating Committee to Helen, enclosing current
materials" (https://www.afb.org/HelenKellerArchive?a=d&d=A-HK01-07-B154-F05-028.1.6).
Helen Keller Archive. American Foundation for the Blind. Retrieved 3 July 2023.
53. "Preparing earth constitution | Global Strategies & Solutions | The Encyclopedia of World
Problems" (https://web.archive.org/web/20230719215501/http://encyclopedia.uia.org/en/stra
tegy/193465). The Encyclopedia of World Problems | Union of International Associations
(UIA). Archived from the original (http://encyclopedia.uia.org/en/strategy/193465) on 19 July
2023. Retrieved 15 July 2023.
Bibliography
Brownlee, George G. (2014). Fred Sanger, double Nobel laureate: a biography. Cambridge,
UK: Cambridge University Press. ISBN 978-1-107-08334-9. Chapters 4-6 contain the 1992
interview that the author conducted with Sanger.
Finch, John (2008), A Nobel Fellow on every floor: a history of the Medical Research
Council Laboratory of Molecular Biology, Cambridge: Medical Research Council, ISBN 978-
1-84046-940-0.
García-Sancho, Miguel (2010). "A new insight into Sanger's development of sequencing:
from proteins to DNA, 1943–1977" (https://www.pure.ed.ac.uk/ws/files/16303515/sangerjhb.
pdf) (PDF). Journal of the History of Biology. 43 (2): 265–323. doi:10.1007/s10739-009-
9184-1 (https://doi.org/10.1007%2Fs10739-009-9184-1). hdl:20.500.11820/e4febe48-772a-
4f47-a1c5-a5ca89505367 (https://hdl.handle.net/20.500.11820%2Fe4febe48-772a-4f47-a1c
5-a5ca89505367). PMID 20665230 (https://pubmed.ncbi.nlm.nih.gov/20665230).
S2CID 1134280 (https://api.semanticscholar.org/CorpusID:1134280).
Sanger, F.; Dowding, M. (1996), Selected Papers of Frederick Sanger: with commentaries,
Singapore: World Scientific, ISBN 978-981-02-2430-1.
Interviews with Nobel Prize–winning scientists: Dr Frederick Sanger (https://www.bbc.co.uk/
archive/scientists/10603.shtml), British Broadcasting Corporation, c. 1985. Interviewed by
Lewis Wolpert. Duration 1 hour.

External links
The Sanger Institute (http://www.sanger.ac.uk/)
About the 1958 Nobel Prize (http://nobelprize.org/nobel_prizes/chemistry/laureates/1958/)
About the 1980 Nobel Prize (http://nobelprize.org/nobel_prizes/chemistry/laureates/1980/)
Fred Sanger (https://web.archive.org/web/20041026014225/http://www.vega.org.uk/series/f
acetoface/sanger/index.php) 2001 Video Documentary by The Vega Science Trust
Portraits of Frederick Sanger (https://www.npg.org.uk/collections/search/person.php?LinkID
=mp06016) at the National Portrait Gallery, London
Frederick Sanger interviewed by Alan Macfarlane, 24 August 2007 (video) (https://www.sms.
cam.ac.uk/media/1130236), also available on Video (https://www.youtube.com/watch?v=0Js
rvWYS7zY) on YouTube. Duration 57 minutes.
Frederick Sanger archive collection (http://wellcomelibrary.org/using-the-library/subject-guid
es/genetics/makers-of-modern-genetics/digitised-archives/fred-sanger/) – Wellcome Library
finding aid for the digitised collection.
Frederick Sanger (https://www.nobelprize.org/laureate/222) on Nobelprize.org

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