Hans Krebs (biochemist)

Sir Hans Adolf Krebs, FRS (/krɛbz, krɛps/, German:

Sir
[hans ˈʔaːdɔlf ˈkʁeːps] ; 25 August 1900 – 22
November 1981)[1][2][3][4] was a German-British
Hans Krebs
FRS
biologist, physician and biochemist.[5] He was a
pioneer scientist in the study of cellular respiration, a
biochemical process in living cells that extracts energy
from food and oxygen and makes it available to drive
the processes of life.[6][7] He is best known for his
discoveries of two important sequences of chemical
reactions that take place in the cells of nearly all
organisms, including humans, other than anaerobic
microorganisms, namely the citric acid cycle and the
urea cycle. The former, often eponymously known as
the "Krebs cycle", is the sequence of metabolic
reactions that allows cells of oxygen-respiring
organisms to obtain far more ATP from the food they
consume than anaerobic processes such as glycolysis
can supply; and its discovery earned Krebs a Nobel
Prize in Physiology or Medicine in 1953. With Hans
Kornberg, he also discovered the glyoxylate cycle,[8] a
Born 25 August 1900
slight variation of the citric acid cycle found in plants,
Hildesheim, Province of Hanover,
bacteria, protists, and fungi.
Kingdom of Prussia, German
Krebs died in 1981 in Oxford, where he had spent 13 Empire
years of his career from 1954 until his retirement in Died 22 November 1981 (aged 81)
1967 at the University of Oxford. Oxford, England, United Kingdom
Citizenship Naturalised British (from 1939)
Education University of Göttingen
Biography University of Freiburg
University of Berlin
University of Hamburg
Early life and education Known for Citric acid cycle
Krebs was born in Hildesheim, Germany, to Georg Urea cycle
Krebs, an ear, nose, and throat surgeon, and Alma Glyoxylate cycle
Krebs (née Davidson).[9] He was of Jewish ancestry Krebs–Henseleit solution
and was the middle of three children.[10] He had an Spouse Margaret Cicely Fieldhouse
elder sister, Elisabeth, and a younger brother, ​
​(m. 1938)​
Wolfgang.[11] Children Paul, John, and Helen
Awards Albert Lasker Award for Basic
Medical Research (1953)
Krebs attended school at the Gymnasium Andreanum Nobel Prize in Physiology or
in his home town.[9] Near the end of World War I, in Medicine (1953)
September 1918, six months short of completing his Royal Medal (1954)
secondary school education, he was conscripted into Copley Medal (1961)
the Imperial German Army.[10] He was allowed to take Scientific career
an emergency examination for his high school
Fields Internal medicine, biochemistry
diploma, which he passed with such a high score that
Institutions Kaiser Wilhelm Institute for
he suspected the examiners of being "unduly lenient
Biology
and sympathetic".[12] With the end of the war two
University of Hamburg
months later, his conscription ended.[10]
University of Cambridge
Krebs decided to follow his father's profession and University of Sheffield
entered the University of Göttingen in December 1918 University of Oxford
to study medicine. In 1919, he transferred to the
University of Freiburg.[9] In 1923, he published his first scientific paper on a tissue staining technique. He
did this work under the guidance of Wilhelm von Mollendorf starting it in 1920.[11] He completed his
medical course in December 1923. To obtain a Doctor of Medicine degree, and a medical license, he
spent one year at the Third Medical Clinic in the University of Berlin.[9] By then he had turned his
professional goal from becoming a practising physician to becoming a medical researcher, particularly in
biochemistry. In 1924, he studied at the Department of Chemistry at the Pathological Institute of the
Charité Hospital, in Berlin, for training in chemistry and biochemistry.[13] He earned his MD degree in
1925 from the University of Hamburg.[10][14][15]

Career
In 1926, Krebs joined Otto Heinrich Warburg as a research assistant at the Kaiser Wilhelm Institute for
Biology in Dahlem, Berlin.[16] He was paid 4800 marks per year. After four years in 1930, with 16
publications to his credit, his mentor Warburg urged him to move on and he took up the position of
Assistant in the Department of Medicine at the Municipal Hospital in Altona (now part of Hamburg). The
next year he moved to the Medical Clinic of the University of Freiburg. At Freiburg, he was in charge of
about 40 patients, and was at liberty to do his own research. Before a year was over at Freiburg, he, with
research student Kurt Henseleit, published their discovery of the ornithine cycle of urea synthesis, which
is the metabolic pathway for urea formation. It is now known as the urea cycle, and is sometimes also
referred to as the Krebs–Henseleit cycle. Together they also developed a complex aqueous solution (a
buffer), or perfusion ex vivo, for studying blood flow in arteries, which is now called the Krebs–Henseleit
buffer.)[17][18] In 1932, he published the basic chemical reactions of the urea cycle, which established his
scientific reputation.

Krebs's life as a respected German scientist came to an abrupt halt in 1933 because of his Jewish ancestry.
With the rise of Hitler's Nazi Party to power, Germany decreed the Law for the Restoration of the
Professional Civil Service, which decreed the removal of all non-Germans, and anti-Nazis, from
professional occupations. Krebs received his official dismissal from his job in April 1933, and his service
was terminated on 1 July 1933. An admirer, Sir Frederick Gowland Hopkins at the University of
Cambridge, immediately came to his rescue, and persuaded the university to recruit Krebs to work with
him in the Department of Biochemistry.[19] By July 1933, he was settled in Cambridge with financial
support from the Rockefeller Foundation.
Although Germany restricted him to bringing only his personal belongings, he was fortunate that the
government agents allowed him to take his equipment and research samples to England. They proved to
be pivotal to his later discoveries, especially the manometer developed by Warburg specifically for the
measurement of oxygen consumption in thin slices of tissues; it was the basis for his research.[20]

He was appointed as Demonstrator in biochemistry in 1934, and in 1935 the University of Sheffield
offered him a post of Lecturer in Pharmacology, with a more spacious laboratory and double the salary.
He worked there for 19 years. The University of Sheffield opened a Department of Biochemistry, now
Department of Molecular Biology and Biotechnology, in 1938 and Krebs became its first Head, and
eventually a Professor in 1945. During his time it became one of the leading departments of biochemistry
in the world.[21] Krebs took over the running of the Sorby Research Institute in 1943. In 1944, the British
Medical Research Council established the MRC Unit for Cell Metabolism Research at Sheffield, and
Krebs was appointed the Director. With this, his laboratory became so large that the locals jokingly
nicknamed it "Krebs's Empire".

He moved with his MRC unit to the University of Oxford in 1954 as Whitley Professor of Biochemistry,
the post he held until his retirement in 1967. The editorial board of Biochemical Journal extended their
good wishes on his retirement, but in return he promised to keep them busy, by producing scientific
papers. He continued his research, and took his MRC unit to the Nuffield Department of Clinical
Medicine at the Radcliffe Infirmary, Oxford. From there he published over 100 research
papers.[14][15][22][23]

Personal life
Krebs met Margaret Cicely Fieldhouse (30 October 1913 – May
1993)[24] when he moved to Sheffield in 1935. They married on
22 March 1938. Krebs later described his life in Sheffield as "19
happy years".[14] They had two sons, Paul (born 1939) and John
(born 1945), and a daughter, Helen (born 1942).[25] John (Sir John
Krebs, and later Baron Krebs) became a renowned ornithologist,
Professor at the University of Oxford, Principal of Jesus College,
Oxford, and Member of the British House of Lords.[26]

Death
After a brief illness, Krebs died on 22 November 1981 in Oxford,
aged 81.[3][27]

Krebs with his wife in Stockholm in Achievements

1953

Urea cycle (Krebs–Henseleit cycle)

In 1932, Krebs worked out the outlines of the urea cycle with a medical student Kurt Henseleit at the
University of Freiburg. While working at the Medical Clinic of the University of Freiburg, Krebs met
Kurt Henseleit, with whom he investigated the chemical process of urea formation. In 1904, two
Germans, A. Kossel and H. D. Dakin, had shown that arginine could be hydrolysed by the enzyme
arginase to form ornithine and urea in inorganic reaction.[28] Based on this reaction, Krebs and Henseleit
postulated that in living cells, similar reaction could occur, and that ornithine and citrulline could be the
intermediate reactions.[29][30] Krebs started working on the possible method for the synthesis of arginine.
Using his Warburg manometer, he mixed a slice of liver with purified ornithine and citrulline. He found
that citrulline acted as a catalyst in the metabolic reactions of urea from ammonia and carbon dioxide. He
and Henseleit published their discovery in 1932. Thus the urea cycle (or "ornithine cycle") was
established, and it was the first metabolic cycle to be discovered.[20][31]

Citric acid cycle (Krebs cycle)

At the University of Sheffield, Krebs and William Arthur Johnson investigated cellular respiration by
which oxygen was consumed to produce energy from the breakdown of glucose. Krebs had earlier
suggested to Warburg while they worked together in Germany that by using a manometer it could be
possible to detect the oxygen consumption and identify the chemical reaction in glucose metabolism.
Warburg had flatly rejected the idea. In Sheffield, Krebs vigorously worked to identify a possible
chemical reaction and came up with numerous hypothetical pathways. Using the manometer he tested
those hypotheses one by one. One hypothesis involving succinate, fumarate, and malate proved to be
useful because all these molecules increased oxygen consumption in the pigeon breast muscle. In 1937,
German biochemists Franz Knoop and Carl Martinus had demonstrated a series of reactions using citrate
that produced oxaloacetate. Krebs realised that these molecules could be the missing intermediates for
such reaction. After four months of experimental works to fill in the gaps, Krebs and Johnson succeeded
in establishing the sequence of the chemical cycle, which they called the "citric acid cycle".[32][33] It is
also known as the "Krebs cycle" or "tricarboxylic acid (TCA) cycle".

Krebs sent a short manuscript account of the discovery to Nature on 10 June 1937. On 14 June, he
received a rejection letter from the editor, saying that the journal had "already sufficient letters to fill
correspondence columns for seven or eight weeks", and encouraging Krebs to "submit it for early
publication to another periodical."[34]

Krebs immediately prepared a longer version titled "The Role of Citric Acid in Intermediate Metabolism
in Animal Tissues", which he sent to the Dutch journal Enzymologia after two weeks and was published
in two months.[3][35] It was followed by a series of papers in different journals.[36][37][38]

Glyoxylate cycle
Krebs continued to add more details to his citric acid cycle. The discovery of acetyl-CoA in 1947 by Fritz
Albert Lipmann was another major contribution.[4][39] However, this new discovery posed a problem in
his classic reaction. In 1957, he, with Hans Kornberg, found that there were additional crucial enzymes.
One was malate synthase, which condenses acetate with glyoxylate to form malate, and the other was
isocitrate lyase, which provides glyoxylate for the reaction by cleaving it from isocitrate.[8] These two
reactions did not follow the normal citric acid cycle, and hence the pathway was named the glyoxylate
bypass of the citric acid cycle, but is now known as the glyoxylate cycle.[20][40]

Honours and awards

Krebs became a naturalised British citizen in 1939. He was
elected Fellow of Trinity College, Oxford, 1954–1967. He was
elected to the Royal Society in 1947. In 1953, he received the
Nobel Prize in Physiology or Medicine for his "discovery of
the citric acid cycle." (He shared the Nobel Prize with Fritz
Lipmann.) For the same reason he was given the Albert Lasker
Award for Basic Medical Research in 1953.[41]

The Royal Society awarded him its Royal Medal in 1954, and
Copley Medal in 1961.[42] He was elected to the American
Academy of Arts and Sciences in 1957.[43]

In 1958, he received the Gold Medal of the Netherlands

Society for Physics, Medical Science and Surgery. He was
knighted in 1958 and was elected Honorary Fellow of Girton
Krebs with Clementine Churchill and
College, Cambridge University, in 1979. He was the Original
Frits Zernike in Stockholm in 1953
Member of the Society for General Microbiology, which
conferred him Honorary Membership in 1980. He was also an
elected member of both the American Philosophical Society and the United States National Academy of
Sciences.[44][45] He received an honorary doctorate from 21 universities.[46]

In July 2015, Krebs's Nobel Prize medal was auctioned off for £225,000 (around $351,225).[47][48] The
proceeds were used to found the Sir Hans Krebs Trust, which provides funding for doctoral students in
the biomedical field and support chemists who had to flee their home countries.[49]

Legacy
The University of Oxford had a building named Hans Krebs Tower, which was occupied by the
Department of Biochemistry. In 2008, a new building for the Department of Biochemistry was
constructed, on which a plaque was placed on 20 May 2013 by the Association of Jewish Refugees.[50]
The plaque was unveiled by John, Lord Krebs, and the inscription reads:[51]

Professor Sir Hans Krebs FRS 1900 – 1981 Biochemist & discoverer of the Krebs cycle Nobel
Prize Winner 1953 worked here 1954 – 1967

The Department of Physiology, Anatomy and Genetics at Oxford University hosts the Sir Hans Krebs
Prize Lecture Series.[52] The last three recipients have been: Jeffrey Friedman M.D., PhD (2019)
Professor Sir Stephen O'Rahilly MD FRS FMedSci (2022) and in 2023 - Professor Dame Frances
Ashcroft FRS FMedSci [53]
The University of Sheffield has The Krebs Institute, founded in 1988. It is a research centre covering
interdisciplinary programmes in biochemical research.[54]

In 1990, the Federation of European Biochemical Societies instituted the Sir Hans Krebs Lecture and
Medal, which was endowed by the Lord Rank Centre for Research. It is awarded for outstanding
achievements in biochemistry and molecular biology.[55][56]

The Society of Friends of Hannover Medical School gives the Sir Hans Krebs Prize, which is worth
10,000 euros.[57][58]

The Biochemical Society offers Krebs Memorial Scholarship to a postgraduate (PhD) student working in
biochemistry or an allied biomedical science at any British university. As of 2014, the scholarship is
worth £18,500 and is given for a year, but is extendable up to three years.[59]

See also
List of Jewish Nobel laureates
List of refugees

References
1. Kornberg, H.; Williamson, D. H. (1984). "Hans Adolf Krebs. 25 August 1900 – 22 November
1981" (https://doi.org/10.1098%2Frsbm.1984.0013). Biographical Memoirs of Fellows of the
Royal Society. 30: 350–385. doi:10.1098/rsbm.1984.0013 (https://doi.org/10.1098%2Frsbm.
1984.0013). PMID 11616005 (https://pubmed.ncbi.nlm.nih.gov/11616005).
2. Weber, George (2001). "Sir Hans A. Krebs Centenary Lecture: cancer and clinical targeting".
Advances in Enzyme Regulation. 41 (1): 1–29. doi:10.1016/S0065-2571(00)00026-1 (http
s://doi.org/10.1016%2FS0065-2571%2800%2900026-1). PMID 11417529 (https://pubmed.n
cbi.nlm.nih.gov/11417529).
3. Gibbons, Marion Stubbs, Geoff (2000). "Hans Adolf Krebs (1900–1981)...His Life and
Times" (https://doi.org/10.1080%2F152165400300001462). IUBMB Life. 50 (3): 163–166.
doi:10.1080/152165400300001462 (https://doi.org/10.1080%2F152165400300001462).
PMID 11142342 (https://pubmed.ncbi.nlm.nih.gov/11142342). S2CID 26393428 (https://api.
semanticscholar.org/CorpusID:26393428).
4. Raju, Tonse NK (1999). "The Nobel Chronicles". The Lancet. 353 (9164): 1628.
doi:10.1016/S0140-6736(05)75758-5 (https://doi.org/10.1016%2FS0140-6736%2805%2975
758-5). PMID 10334294 (https://pubmed.ncbi.nlm.nih.gov/10334294). S2CID 54354255 (htt
ps://api.semanticscholar.org/CorpusID:54354255).
5. "Sir Hans Adolf Krebs" (http://www.britannica.com/nobelprize/article-9046221?tocId=904622
1). Encyclopædia Britannica, Inc. Retrieved 26 July 2014.
6. Leigh, F W (2009). "Sir Hans Adolf Krebs (1900–81), pioneer of modern medicine, architect
of intermediary metabolism". Journal of Medical Biography. 17 (3): 149–154.
doi:10.1258/jmb.2009.009032 (https://doi.org/10.1258%2Fjmb.2009.009032).
PMID 19723965 (https://pubmed.ncbi.nlm.nih.gov/19723965). S2CID 207200346 (https://ap
i.semanticscholar.org/CorpusID:207200346).
7. Edsall, John T (1993). "Master of metabolic cycles". Nature. 366 (6454): 417–418.
Bibcode:1993Natur.366..417E (https://ui.adsabs.harvard.edu/abs/1993Natur.366..417E).
doi:10.1038/366417a0 (https://doi.org/10.1038%2F366417a0). S2CID 4241345 (https://api.s
emanticscholar.org/CorpusID:4241345).
 8. Kornberg, HL; Krebs, HA (1957). "Synthesis of cell constituents from C2-units by a modified
tricarboxylic acid cycle". Nature. 179 (4568): 988–91. Bibcode:1957Natur.179..988K (https://
ui.adsabs.harvard.edu/abs/1957Natur.179..988K). doi:10.1038/179988a0 (https://doi.org/10.
1038%2F179988a0). PMID 13430766 (https://pubmed.ncbi.nlm.nih.gov/13430766).
S2CID 40858130 (https://api.semanticscholar.org/CorpusID:40858130).
9. "The Nobel Prize in Physiology or Medicine 1953" (https://www.nobelprize.org/prizes/medici
ne/1953/krebs/biographical/). NobelPrize.org. 1953. Retrieved 11 June 2024.
10. Nicholls, Mark (14 September 2020). "Hans Krebs". European Heart Journal. 41 (35): 3294–
3296. doi:10.1093/eurheartj/ehaa604 (https://doi.org/10.1093%2Feurheartj%2Fehaa604).
ISSN 0195-668X (https://search.worldcat.org/issn/0195-668X). PMID 33033838 (https://pub
med.ncbi.nlm.nih.gov/33033838).
11. Roth, Klaus (3 November 2020). "Sir Hans Adolf Krebs (1900 – 1981) – Part 1".
ChemViews. doi:10.1002/chemv.202000103 (https://doi.org/10.1002%2Fchemv.20200010
3).
12. Hans, Krebs; Martin, Anne (1981). Reminiscences and Reflections (https://books.google.co
m/books?id=27-4AAAAIAAJ). Oxford: Clarendon Press. p. 14. ISBN 978-0-1985-4702-0.
13. Holmes, Frederic Lawrence (1991). Hans Krebs (https://books.google.com/books?id=qi8qfn
KTuGoC&pg=PA121). Monographs on the History and Philosophy of Biology. Oxford
University Press. p. 121. ISBN 978-0-19-536128-5. Retrieved 12 June 2024.
14. Willcocks, Elizabeth (1 June 2003). "A lucky man" (http://www.rsc.org/chemistryworld/Issue
s/2003/June/luckyman.asp). Royal Society of Chemistry.
15. Quayle, J. R. (1982). "Obituary" (https://doi.org/10.1099%2F00221287-128-10-2215).
Microbiology. 128 (10): 2215–2220. doi:10.1099/00221287-128-10-2215 (https://doi.org/10.
1099%2F00221287-128-10-2215). PMID 6759612 (https://pubmed.ncbi.nlm.nih.gov/675961
2).
16. Krebs, Hans A.; Lipmann, Fritz (1974). "Dahlem in the Late Nineteen Twenties" (https://book
s.google.com/books?id=kaTiDwAAQBAJ&pg=PA7). In Richter, Dietmar (ed.). Lipmann
Symposium. Energy transformation in biological systems. Berlin: Walter De Gruyter. pp. 7–
27. ISBN 9783112316269.
17. Bailey, Leslie E; Ong, Seok D (1978). "Krebs–Henseleit solution as a physiological buffer in
perfused and superfused preparations". Journal of Pharmacological Methods. 1 (2): 171–
175. doi:10.1016/0160-5402(78)90022-0 (https://doi.org/10.1016%2F0160-5402%2878%29
90022-0).
18. "Krebs–Henseleit Buffer Modified" (http://www.sigmaaldrich.com/catalog/product/sigma/k375
3?lang=en&region=IN). Sigma-Aldrich Co. LLC. Retrieved 28 July 2014.
19. Hans, Krebs; Martin, Anne (1981). Reminiscences and Reflections (https://books.google.co
m/books?id=27-4AAAAIAAJ). Oxford: Clarendon Press. p. 48. ISBN 978-0-1985-4702-0.
20. Wilson, B. A.; Schisler, J. C.; Willis, M. S. (2010). "Sir Hans Adolf Krebs: Architect of
Metabolic Cycles" (https://doi.org/10.1309%2FLMZ5ZLAC85GFMGHU). Laboratory
Medicine. 41 (6): 377–380. doi:10.1309/LMZ5ZLAC85GFMGHU (https://doi.org/10.1309%2
FLMZ5ZLAC85GFMGHU).
21. Among the distinguished biochemists who worked at Sheffield during that period were Keith
Dalziel, Quentin Gibson, Hans Kornberg, Vincent Massey and Gregorio Weber
22. Williamson, DH (1982). "Sir Hans Krebs (1900–1981)" (https://www.ncbi.nlm.nih.gov/pmc/art
icles/PMC1158308). The Biochemical Journal. 204 (1): 1–2. doi:10.1042/bj2040001 (https://
doi.org/10.1042%2Fbj2040001). PMC 1158308 (https://www.ncbi.nlm.nih.gov/pmc/articles/P
MC1158308). PMID 7052063 (https://pubmed.ncbi.nlm.nih.gov/7052063).
23. Williamson, DH (1981). "Tribute to Sir Hans Krebs". Biochemical Society Transactions. 9 (1):
1–2. doi:10.1042/bst0090001 (https://doi.org/10.1042%2Fbst0090001). PMID 7011867 (http
s://pubmed.ncbi.nlm.nih.gov/7011867).
24. "Margaret Cicely Krebs" (https://www.geni.com/people/Margaret-Krebs/6000000029297312
586). 22 June 2018.
25. "Lady Margaret Krebs" (https://web.archive.org/web/20140809145137/http://www.wickersley
web.co.uk/hist/krebs.htm). wickersleyweb.co.uk. Archived from the original (http://www.wick
ersleyweb.co.uk/hist/krebs.htm) on 9 August 2014. Retrieved 29 July 2014.
26. "Lord Krebs | Jesus College, University of Oxford" (https://web.archive.org/web/2011100715
1631/http://www.jesus.ox.ac.uk/fellows-and-staff/fellows/lord-krebs). 7 October 2011.
Archived from the original (http://www.jesus.ox.ac.uk/fellows-and-staff/fellows/lord-krebs) on
7 October 2011. Retrieved 13 March 2024.
27. Altman, Lawrence K. (9 December 1981). "Sir Hans Krebs, winner of Nobel for research on
food cycles, dies" (https://www.nytimes.com/1981/12/09/obituaries/sir-hans-krebs-winner-of-
nobel-for-research-on-food-cycles-dies.html). The New York Times. Retrieved 13 December
2014.
28. Kinne-Saffran, E; Kinne, RK (1999). "Vitalism and synthesis of urea. From Friedrich Wöhler
to Hans A. Krebs". American Journal of Nephrology. 19 (2): 290–294.
doi:10.1159/000013463 (https://doi.org/10.1159%2F000013463). PMID 10213830 (https://p
ubmed.ncbi.nlm.nih.gov/10213830). S2CID 71727190 (https://api.semanticscholar.org/Corp
usID:71727190).
29. Graßhoff, Gerd; May, Michael (2003). "Hans Krebs' and Kurt Henseleit's Laboratory
Notebooks and Their Discovery of the Urea Cycle-Reconstructed with Computer Models". In
Holmes, Frederic L.; Renn, Jürgen; Rheinberger, Hans-Jörg (eds.). Reworking the Bench :
Research Notebooks in the History of Science. Dordrecht: Kluwer Academic Publishers.
pp. 269–294. doi:10.1007/0-306-48152-9_13 (https://doi.org/10.1007%2F0-306-48152-9_1
3). ISBN 978-1-4020-1039-2.
30. Nickelsen, Kärin; Graßhoff, Gerd (2009). Hon, Giora; Schickore, Jutta; Steinle, Friedrich
(eds.). Going Amiss in Experimental Research. Dordrect, Netherlands: Springer Verlag.
pp. 91–117. doi:10.1007/978-1-4020-8893-3_7 (https://doi.org/10.1007%2F978-1-4020-889
3-3_7). ISBN 978-1-4020-8892-6.
31. Holmes, FL (1980). "Hans Krebs and the discovery of the ornithine cycle". Federation
Proceedings. 39 (2): 216–25. PMID 6986292 (https://pubmed.ncbi.nlm.nih.gov/6986292).
32. Krebs, HA; Johnson, WA (1937). "Metabolism of ketonic acids in animal tissues" (https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC1266984). The Biochemical Journal. 31 (4): 645–60.
doi:10.1042/bj0310645 (https://doi.org/10.1042%2Fbj0310645). PMC 1266984 (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC1266984). PMID 16746382 (https://pubmed.ncbi.nlm.nih.
gov/16746382).
33. Krebs, HA; Johnson, WA (1937). "Acetopyruvic acid (alphagamma-diketovaleric acid) as an
intermediate metabolite in animal tissues" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1
267003). The Biochemical Journal. 31 (5): 772–9. doi:10.1042/bj0310772 (https://doi.org/10.
1042%2Fbj0310772). PMC 1267003 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC12670
03). PMID 16746397 (https://pubmed.ncbi.nlm.nih.gov/16746397).
34. Borrel, Brendan (1 March 2010). "Nature rejects Krebs's paper, 1937" (http://www.the-scienti
st.com/?articles.view/articleNo/28819/title/Nature-rejects-Krebs-s-paper--1937/). The
Scientist. Retrieved 29 July 2014.
35. Lalchhandama, K (2012). "How to write a rather tenacious and legible scientific paper" (http
s://web.archive.org/web/20140427025145/http://www.sciencevision.org/current_issue/dl/Lal
chhandama.pdf) (PDF). Science Vision. 12 (2): 41–54. Archived from the original (http://ww
w.sciencevision.org/current_issue/dl/Lalchhandama.pdf) (PDF) on 27 April 2014.
36. Krebs, H.A. (1937). "The Intermediate Metabolism of Carbohydrates". The Lancet. 230
(5952): 736–738. doi:10.1016/S0140-6736(00)88690-0 (https://doi.org/10.1016%2FS0140-6
736%2800%2988690-0).
37. Krebs, HA; Salvin, E; Johnson, WA (1938). "The formation of citric and alpha-ketoglutaric
acids in the mammalian body" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1264001).
The Biochemical Journal. 32 (1): 113–7. doi:10.1042/bj0320113 (https://doi.org/10.1042%2F
bj0320113). PMC 1264001 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1264001).
PMID 16746585 (https://pubmed.ncbi.nlm.nih.gov/16746585).
38. Krebs, HA (1938). "Micro-determination of alpha-ketoglutaric acid" (https://www.ncbi.nlm.nih.
gov/pmc/articles/PMC1264000). The Biochemical Journal. 32 (1): 108–12.
doi:10.1042/bj0320108 (https://doi.org/10.1042%2Fbj0320108). PMC 1264000 (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC1264000). PMID 16746584 (https://pubmed.ncbi.nlm.nih.
gov/16746584).
39. Buchanan, JM (2002). "Biochemistry during the life and times of Hans Krebs and Fritz
Lipmann" (https://doi.org/10.1074%2Fjbc.R200019200). The Journal of Biological
Chemistry. 277 (37): 33531–6. doi:10.1074/jbc.R200019200 (https://doi.org/10.1074%2Fjbc.
R200019200). PMID 12070179 (https://pubmed.ncbi.nlm.nih.gov/12070179).
40. Kornberg, Hans (2000). "Krebs and his trinity of cycles". Nature Reviews Molecular Cell
Biology. 1 (3): 225–228. doi:10.1038/35043073 (https://doi.org/10.1038%2F35043073).
PMID 11252898 (https://pubmed.ncbi.nlm.nih.gov/11252898). S2CID 28092593 (https://api.
semanticscholar.org/CorpusID:28092593).
41. "Albert Lasker Basic Medical Research Award" (https://web.archive.org/web/201410140002
29/http://www.laskerfoundation.org/awards/1953basic.htm). Lasker Foundation. Archived
from the original (http://www.laskerfoundation.org/awards/1953basic.htm) on 14 October
2014. Retrieved 29 July 2014.
42. Hans Krebs' CV (https://www.leopoldina.org/fileadmin/redaktion/Mitglieder/CV_Krebs_Hans
_D.pdf), Leopoldina.org. Accessed 8 February 2023.
43. "Hans Adolf Krebs" (https://www.amacad.org/person/hans-adolf-krebs). American Academy
of Arts & Sciences. Retrieved 29 November 2022.
44. "APS Member History" (https://search.amphilsoc.org/memhist/search?creator=Hans+Krebs
&title=&subject=&subdiv=&mem=&year=&year-max=&dead=&keyword=&smode=advance
d). search.amphilsoc.org. Retrieved 29 November 2022.
45. "Hans Krebs" (http://www.nasonline.org/member-directory/deceased-members/20001329.ht
ml). www.nasonline.org. Retrieved 29 November 2022.
46. Kronberg, Hans (2004). "Krebs, Sir Hans Adolf (1900–1981), biochemist". Oxford Dictionary
of National Biography. Vol. 1 (online ed.). Oxford University Press.
doi:10.1093/ref:odnb/31327 (https://doi.org/10.1093%2Fref%3Aodnb%2F31327).
(Subscription or UK public library membership (https://www.oxforddnb.com/help/subscribe#publi
c) required.)
47. "Sheffield academic's Nobel Prize medal sells for £225,000" (https://www.bbc.com/news/uk-
england-south-yorkshire-33522526). BBC. 14 July 2015.
48. Keener, Amanda B. (16 July 2015). "Krebs Nobel Auctioned" (http://www.the-scientist.com/?
articles.view/articleNo/43521/title/Krebs-Nobel-Auctioned). TheScientist.
49. "Sir Hans Krebs Trust / Cara Fellowships" (https://www.cara.ngo/sir-hans-krebs-trust-cara-fe
llowships/). Cara. 1 December 2015. Retrieved 19 November 2019.
50. Shock, Kathy (2013). "New Blue Plaque in Oxford" (https://web.archive.org/web/201408091
25740/http://www.oxfordjewishheritage.co.uk/news-a-events/news/221-new-blue-plaque-in-
oxford). Oxford Jewish Heritage. Archived from the original (http://www.oxfordjewishheritag
e.co.uk/news-a-events/news/221-new-blue-plaque-in-oxford) on 9 August 2014. Retrieved
28 July 2014.
51. Itzhaki, Jane (22 May 2013). "Plaque unveiled to honour work of Sir Hans Krebs" (https://we
b.archive.org/web/20140812215942/http://www.bioch.ox.ac.uk/aspsite/index.asp?pageid=98
1). Department of Biochemistry, University of Oxford. Archived from the original (http://www.
bioch.ox.ac.uk/aspsite/index.asp?pageid=981) on 12 August 2014. Retrieved 28 July 2014.
52. "Sir Hans Krebs Prize Lecture Series" (https://www.dpag.ox.ac.uk/named-lectures/hans-kre
bs-lecture).
53. "About []Sir Hans Krebs Prize Lecture Series" (https://www.dpag.ox.ac.uk/named-lectures/h
ans-krebs-lecture).
54. "About [The Krebs Institute]" (https://web.archive.org/web/20150510025821/http://www.kreb
s.group.shef.ac.uk/AboutKI.html). The Krebs Institute. Archived from the original (http://www.
krebs.group.shef.ac.uk/AboutKI.html) on 10 May 2015. Retrieved 28 July 2014.
55. "FEBS Medals" (http://www.febs.org/our-activities/awards/medals). Federation of European
Biochemical Societies. Retrieved 28 July 2014.
56. "The Sir Hans Krebs Medal of the Federation of European Biochemical Societies" (http://ww
w.lifesci.dundee.ac.uk/awards/sir-hans-krebs-medal-federation-european-biochemical-societ
ies). University of Dundee. 20 August 2013. Retrieved 28 July 2014.
57. "Heineke receives the Sir Hans Krebs Prize" (http://www.rebirth-hannover.de/en/press-medi
a/press-releases/singleview.html?tx_ttnews%5byear%5d=2011&tx_ttnews%5bmonth%5d=1
1&tx_ttnews%5bday%5d=02&tx_ttnews%5btt_news%5d=27&cHash=1213fd16c777c3bd44
dfc17c2d8dc36e). Rebirth Hannover. 2 December 2011. Retrieved 29 July 2014.
58. "Ivan Dikic is awarded the Sir Hans Krebs Prize for his research on ubiquitin" (https://web.ar
chive.org/web/20180720140005/https://www.bifonds.de/news-network/news-pictures/news-i
tem/items/ivan-dikic-is-awarded-the-sir-hans-krebs-prize-for-his-research-on-ubiquitin.html).
Boehringer Ingelheim Fonds. 13 November 2009. Archived from the original (http://www.bifo
nds.de/news-network/news-pictures/news-item/items/ivan-dikic-is-awarded-the-sir-hans-kre
bs-prize-for-his-research-on-ubiquitin.html) on 20 July 2018. Retrieved 29 July 2014.
59. "Krebs Memorial Scholarship" (https://web.archive.org/web/20140729165412/http://www.bio
chemistry.org/Grants/KrebsMemorialScholarship.aspx). Biochemical Society. Archived from
the original (http://www.biochemistry.org/Grants/KrebsMemorialScholarship.aspx) on 29 July
2014. Retrieved 29 July 2014.

Bibliography
Medawar, Jean; Pyke, David (2012). Hitler's Gift: The True Story of the Scientists Expelled
by the Nazi Regime (Paperback). New York: Arcade Publishing. ISBN 978-1-61145-709-4.
Oakes, Elizabeth H (2007). Encyclopedia of World Scientists (https://books.google.com/boo
ks?id=uPRB-OED1bcC) (Rev. ed.). New York: Facts on File. p. 414. ISBN 978-1-4381-
1882-6.
Holmes, Frederic Lawrence (1991). Hans Krebs : Volume 1: The Formation of a Scientific
Life, 1900–1933 (https://books.google.com/books?id=qi8qfnKTuGoC). New York: Oxford
University Press. p. 512. ISBN 978-0-195-07072-9.

External links
Hans Krebs (https://www.nobelprize.org/laureate/354) on Nobelprize.org including the Nobel
Lecture, 11 December 1953 The Citric Acid Cycle
1953 Prize in Physiology or Medicine (http://nobelprize.org/nobel_prizes/medicine/laureates/
1953/) Nobel Prizes
Deconstructing the Tour, University of Sheffield (biography) (http://www.deconstructingthetou
r.group.shef.ac.uk/krebs-cycle/) Archived (https://web.archive.org/web/20210419085915/htt
p://www.deconstructingthetour.group.shef.ac.uk/krebs-cycle/) 19 April 2021 at the Wayback
Machine
Sir Hans Adolf Krebs at whonamedit (http://www.whonamedit.com/doctor.cfm/1541.html)
 Catalogue of the papers and correspondence of Sir Hans Adolf Krebs, FRS (1900–1981) (ht
tp://apps.nationalarchives.gov.uk/a2a/records.aspx?cat=200-csac113486_3&cid=0#0)
Hans A. Krebs (https://nobel.bh.org.il/en/persona/Hans-Krebs/), Nobel Luminaries – Jewish
Nobel Prize Winners, on the Beit Hatfutsot-The Museum of the Jewish People (https://www.
bh.org.il/) Website.

Retrieved from "https://en.wikipedia.org/w/index.php?title=Hans_Krebs_(biochemist)&oldid=1259847536"