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While on the faculty of the [[University of Wisconsin]], he shared the 1968 [[Nobel Prize for Physiology or Medicine|Nobel Prize]] with [[Marshall W. Nirenberg]] and [[Robert W. Holley]] for researchthat showed how the order of [[nucleotides]] in [[nucleic acids]], which carry the [[genetic code]] of the cell, control the cell’s synthesis of proteins. Khorana and Nirenberg were also awarded the [[Louisa Gross Horwitz Prize]] from [[Columbia University]] in the same year.<ref>{{cite web |url=http://www.cumc.columbia.edu/horwitz |title=The Official Site of Louisa Gross Horwitz Prize}}</ref><ref>{{cite journal |author=Thomas P. Sakmar |title=Har Gobind Khorana (1922–2011): Pioneering Spirit |year=2012 |journal=[[PLoS Biology]] |volume=10 |issue=2 |pages= e1001273|doi=10.1371/journal.pbio.1001273}}</ref>
While on the faculty of the [[University of Wisconsin]], he shared the 1968 [[Nobel Prize for Physiology or Medicine|Nobel Prize]] with [[Marshall W. Nirenberg]] and [[Robert W. Holley]] for researchthat showed how the order of [[nucleotides]] in [[nucleic acids]], which carry the [[genetic code]] of the cell, control the cell’s synthesis of proteins. Khorana and Nirenberg were also awarded the [[Louisa Gross Horwitz Prize]] from [[Columbia University]] in the same year.<ref>{{cite web |url=http://www.cumc.columbia.edu/horwitz |title=The Official Site of Louisa Gross Horwitz Prize}}</ref><ref>{{cite journal |author=Thomas P. Sakmar |title=Har Gobind Khorana (1922–2011): Pioneering Spirit |year=2012 |journal=[[PLoS Biology]] |volume=10 |issue=2 |pages= e1001273|doi=10.1371/journal.pbio.1001273}}</ref>


Khorana was born in [[Raipur]], India, (today [[Kabirwala]] in [[Pakistan]]). He was the youngest of five children of Ganpat Rai Khorana, a taxation clerk, and Krishna Devi Khorana. He served on the faculty of three universities in North America. He became a [[naturalized citizen]] of the [[United States]] in 1966,<ref name="brit">{{cite web |url=http://www.britannica.com/EBchecked/topic/316846/Har-Gobind-Khorana |title=HG Khorana Britannica}}</ref> and subsequently received the [[National Medal of Science]] in 1987.<ref>{{cite web|url=https://www.usatoday.com/story/tech/talkingtech/2018/01/09/google-doodle-honors-dna-researcher-har-gobind-khorana/1016000001/|title=Google Doodle honors DNA researcher Har Gobind Khorana|website=usatoday.com|accessdate=9 January 2018}}</ref> He co-directed the Institute for Enzyme Research,<ref name="news.wisc.edu">{{cite web |url=http://news.wisc.edu/biochemist-har-gobind-khorana-whose-uw-work-earned-the-nobel-prize-dies/ |title=Biochemist Har Gobind Khorana, whose UW work earned the Nobel Prize, dies |website=news.wisc.edu |access-date=2017-01-10}}</ref> became a professor of biochemistry in 1962 and was named Conrad A. Elvehjem Professor of Life Sciences at [[University of Wisconsin–Madison]].<ref>{{cite journal |last1=Kresge |first1=Nicole |last2=Simoni |first2=Robert D. |last3=Hill |first3=Robert L. |date=2009-05-29 |title=Total Synthesis of a Tyrosine Suppressor tRNA: the Work of H. Gobind Khorana |journal=The Journal of Biological Chemistry |volume=284 |issue=22 |pages=e5 |issn=0021-9258 |pmc=2685647}}</ref> He served as [[Massachusetts Institute of Technology|MIT]]'s Alfred P. Sloan Professor of Biology and Chemistry, Emeritus<ref>{{Cite news|url=http://news.mit.edu/2011/obit-khorana-1110|title=Gobind Khorana, MIT professor emeritus, dies at 89|work=MIT News|access-date=2018-01-09}}</ref><ref>{{cite web |url=http://web.mit.edu/chemistry/www/faculty/khorana.html |title=MIT HG Khorana MIT laboratory}}</ref> and was a member of the Board of Scientific Governors at [[The Scripps Research Institute]].
Khorana was born in [[Raipur]], India. He was the youngest of five children of Ganpat Rai Khorana, a taxation clerk, and Krishna Devi Khorana. He served on the faculty of three universities in North America. He became a [[naturalized citizen]] of the [[United States]] in 1966,<ref name="brit">{{cite web |url=http://www.britannica.com/EBchecked/topic/316846/Har-Gobind-Khorana |title=HG Khorana Britannica}}</ref> and subsequently received the [[National Medal of Science]] in 1987.<ref>{{cite web|url=https://www.usatoday.com/story/tech/talkingtech/2018/01/09/google-doodle-honors-dna-researcher-har-gobind-khorana/1016000001/|title=Google Doodle honors DNA researcher Har Gobind Khorana|website=usatoday.com|accessdate=9 January 2018}}</ref> He co-directed the Institute for Enzyme Research,<ref name="news.wisc.edu">{{cite web |url=http://news.wisc.edu/biochemist-har-gobind-khorana-whose-uw-work-earned-the-nobel-prize-dies/ |title=Biochemist Har Gobind Khorana, whose UW work earned the Nobel Prize, dies |website=news.wisc.edu |access-date=2017-01-10}}</ref> became a professor of biochemistry in 1962 and was named Conrad A. Elvehjem Professor of Life Sciences at [[University of Wisconsin–Madison]].<ref>{{cite journal |last1=Kresge |first1=Nicole |last2=Simoni |first2=Robert D. |last3=Hill |first3=Robert L. |date=2009-05-29 |title=Total Synthesis of a Tyrosine Suppressor tRNA: the Work of H. Gobind Khorana |journal=The Journal of Biological Chemistry |volume=284 |issue=22 |pages=e5 |issn=0021-9258 |pmc=2685647}}</ref> He served as [[Massachusetts Institute of Technology|MIT]]'s Alfred P. Sloan Professor of Biology and Chemistry, Emeritus<ref>{{Cite news|url=http://news.mit.edu/2011/obit-khorana-1110|title=Gobind Khorana, MIT professor emeritus, dies at 89|work=MIT News|access-date=2018-01-09}}</ref><ref>{{cite web |url=http://web.mit.edu/chemistry/www/faculty/khorana.html |title=MIT HG Khorana MIT laboratory}}</ref> and was a member of the Board of Scientific Governors at [[The Scripps Research Institute]].


==Biography==
==Biography==

Revision as of 21:56, 9 January 2018

Har Gobind Khorana
Born(1922-01-09)January 9, 1922
Raipur, Punjab, British India, Present day India
DiedNovember 9, 2011(2011-11-09) (aged 89)
Concord, MA
CitizenshipAmerican
Alma mater
Known forFirst to demonstrate the role of nucleotides in protein synthesis
Awards
Scientific career
FieldsMolecular biology
Institutions
Doctoral advisorRoger J.S. Beer
Doctoral studentsShiladitya DasSarma
Signature
Har Gobind Khorana signature

Har Gobind Khorana (January 9, 1922 – November 9, 2011)[1][2][3] was an Indian American[4] biochemist.

While on the faculty of the University of Wisconsin, he shared the 1968 Nobel Prize with Marshall W. Nirenberg and Robert W. Holley for researchthat showed how the order of nucleotides in nucleic acids, which carry the genetic code of the cell, control the cell’s synthesis of proteins. Khorana and Nirenberg were also awarded the Louisa Gross Horwitz Prize from Columbia University in the same year.[5][6]

Khorana was born in Raipur, India. He was the youngest of five children of Ganpat Rai Khorana, a taxation clerk, and Krishna Devi Khorana. He served on the faculty of three universities in North America. He became a naturalized citizen of the United States in 1966,[7] and subsequently received the National Medal of Science in 1987.[8] He co-directed the Institute for Enzyme Research,[9] became a professor of biochemistry in 1962 and was named Conrad A. Elvehjem Professor of Life Sciences at University of Wisconsin–Madison.[10] He served as MIT's Alfred P. Sloan Professor of Biology and Chemistry, Emeritus[11][12] and was a member of the Board of Scientific Governors at The Scripps Research Institute.

Biography

Har Gobind Khorana was born to parents Shrimati Krishna Khorana Devi, and Shri Ganpat Rai Khorana in Raipur, a village in Punjab, which is now part of India.[13] The exact date of his birth is not certain but he believed that it might have been January 9, 1922;[14] this date was later shown in some documents, and has been widely accepted.[15] He was the youngest of a family that included one daughter and four sons. His father was a patwari, a village agricultural taxation clerk in the Indian government. In his autobiography, Khorana wrote this summary. "Although poor, my father was dedicated to educating his children and we were practically the only literate family in the village inhabited by about 100 people."[16] The first four years of his education were provided under a tree, a spot that was, in effect, the only school in the village.[13]

Har Gobind Khorana attended D.A.V. High School in Multan, West Punjab.[13] Later, he studied at the Punjab University in Lahore, with the assistance of scholarships, where he obtained a Bachelor's degree in 1943[17] and an M. Sc. degree in 1948.[18][19]

Khorana lived in India until 1945, when the award of a Government of India Fellowship made it possible for him to go to England[20] where he studied for, and received in 1948, a Ph. D. degree at the University of Liverpool in organic chemistry, working with Roger S. Beer.[21][22][16] Khorana spent a postdoctoral year (1948-1949) at the Eidgenössische Technische Hochschule in Zurich, Switzerland with Professor Vladimir Prelog.[16] He worked for nearly a year on alkaloid chemistry in an unpaid position.[23][13]

During a brief period in 1949, he was unable to find a job in his home province of the Punjab, which was by then part of India.[13] In the fall of that year, Khorana returned to England where he obtained a fellowship to work with G. W. Kenner and A. R. Todd on peptides and nucleotides.[23] He stayed in Cambridge from 1950 until 1952.

After another postdoctoral year at Cambridge University, he accepted a job offer in 1952, from the British Columbia Research Council in Canada at University of British Columbia[24] and the family settled in Vancouver, British Columbia.[17] Khorana was excited by the prospect of starting his own lab, a colleague later recalled.[25]His mentor said that the Council had few facilities at the time but gave the researcher "all the freedom in the world".[26] His work in British Columbia was on "nucleic acids and synthesis of many important biomolecules" according to the American Chemical Society.[27]

In 1960 Khorana moved to the Institute for Enzyme Research at the University of Wisconsin at Madison as co-director of the Institute for Enzyme Research.[28] He became a US citizen in 1966.[29] Starting in the fall of 1970, Khorana was the Alfred P. Sloan Professor of Biology and Chemistry at the Massachusetts Institute of Technology.[30] While at Wisconsin, "he helped decipher the mechanisms by which RNA codes for the synthesis of proteins" and "began to work on synthesizing functional genes" according to the American Chemical Society.[31] He retired from MIT in 2007.[29]

Har Gobind Khorana was married in 1952 to Esther Elizabeth Sibler, of Swiss origin. They had met in Switzerland and had three children, Julia Elizabeth, Emily Anne and Dave Roy.

Research

Ribonucleic acid (RNA) with three repeating units (UCUCUCU → UCU CUC UCU) produced two alternating amino acids. This, combined with the Nirenberg and Leder experiment, showed that UCU genetically codes for serine and CUC codes for leucine. RNAs with three repeating units (UACUACUA → UAC UAC UAC, or ACU ACU ACU, or CUA CUA CUA) produced three different strings of amino acids. RNAs with four repeating units including UAG, UAA, or UGA, produced only dipeptides and tripeptides thus revealing that UAG, UAA and UGA are stop codons.[32]

Their Nobel lecture was delivered on 12 December 1968.[32] Khorana was the first scientist to chemically synthesize oligonucleotides.[33] This achievement, in the 1970s, was also the world's first synthetic gene; in later years, the process has become widespread.[30] Subsequent scientists referred to his research while advancing genome editing with the CRISPR/Cas9 system.[29]

Subsequent research

He extended the above to long DNA polymers using non-aqueous chemistry and assembled these into the first synthetic gene, using polymerase and ligase enzymes that link pieces of DNA together,[33] as well as methods that anticipated the invention of polymerase chain reaction (PCR).[34] These custom-designed pieces of artificial genes are widely used in biology labs for sequencing, cloning and engineering new plants and animals, and are integral to the expanding use of DNA analysis to understand gene-based human disease as well as human evolution. Khorana's invention(s) have become automated and commercialized so that anyone now can order a synthetic oligonucleotide or a gene from any of a number of companies. One merely needs to send the genetic sequence to one of the companies to receive an oligonucleotide with the desired sequence.

After the middle of the 1970s, his lab studied the biochemistry of bacteriorhodopsin, a membrane protein that converts light energy into chemical energy by creating a proton gradient.[35] Later, his lab went on to study the structurally related visual pigment known as rhodopsin.[36]

A summary of his work was provided by a former colleague at the University of Wisconsin. "Khorana was an early practitioner, and perhaps a founding father, of the field of chemical biology. He brought the power of chemical synthesis to bear on deciphering the genetic code, relying on different combinations of trinucleotides."[37][38]

Awards and honors

In addition to sharing the Nobel prize (while he was working at the University of Wisconsin in the U.S.),[18] Khorana was elected as Foreign Member of the Royal Society (ForMemRS) in 1978.[39] The University of Wisconsin-Madison, the Government of India (DBT Department of Biotechnology), and the Indo-US Science and Technology Forum, in 2007, created the Khorana Program, jointly. The mission of the Khorana Program is to build a seamless community of scientists, industrialists, and social entrepreneurs in the United States and India.

The program is focused on three objectives:[40] Providing graduate and undergraduate students with a transformative research experience, engaging partners in rural development and food security, and facilitating public-private partnerships between the U.S. and India. In 2009, Khorana was hosted by the Khorana Program and honored at the 33rd Steenbock Symposium in Madison, Wisconsin.[9]

Other honours included the Louisa Gross Horwitz Prize from Columbia University and the Lasker Foundation Award for Basic Medical Research, both in 1969, the Willard Gibbs Medal of the Chicago section of the American Chemical Society, in 1974, the Gairdner Foundation Annual Award, in 1980 and the Paul Kayser International Award of Merit in Retina Research, in 1987.[16]

On 9 January 2018, the worldwide Google network paid homage to the researcher, citing his Nobel prize award and National Medal of Science (1987). The latter was presented for "his innovative contributions that significantly contributed to our understanding of gene structure, membrane function and vision."[41] The illustration of Har Gobind Khorana's likeness depicted his use of three-letter combinations to represent the four chemical bases found in RNA.[42] According to the Washington Post, "Dr. Khorana confirmed the belief that the code is transmitted by separate and distinct three-letter “words” and found the precise order of the nucleotides within each triplet".[43]

Death

Har Gobind Khorana died on November 9, 2011 in Concord, Massachusetts, at the age of 89.[44] His wife Esther and daughter Emily Anne had died earlier[45] but Khorana was survived by his other two children.[14] Julia later wrote about his work as a professor. "Even while doing all this research, he was always really interested in education, in students and young people."[16]

In his obituary, the Washington Post provided this summary of the man: "Dr. Khorana was known for a modest, ingratiating manner. He tended to shun publicity, making many of his most important scientific announcements at departmental seminars and in scientific publications".[46]

References

  1. ^ Caruthers, M.; Wells, R. (2011). "Har Gobind Khorana (1922-2011)". Science. 334 (6062): 1511. Bibcode:2011Sci...334.1511C. doi:10.1126/science.1217138. PMID 22174242.
  2. ^ Rajbhandary, U. L. (2011). "Har Gobind Khorana (1922–2011)". Nature. 480 (7377): 322. Bibcode:2011Natur.480..322R. doi:10.1038/480322a. PMID 22170673.
  3. ^ Gellene, Denise (2011-11-14). "H. Gobind Khorana, 1968 Nobel Winner for RNA Research, Dies". The New York Times. ISSN 0362-4331. Retrieved 2018-01-09.
  4. ^ "Har Gobind Khorana Biography".
  5. ^ "The Official Site of Louisa Gross Horwitz Prize".
  6. ^ Thomas P. Sakmar (2012). "Har Gobind Khorana (1922–2011): Pioneering Spirit". PLoS Biology. 10 (2): e1001273. doi:10.1371/journal.pbio.1001273.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  7. ^ "HG Khorana Britannica".
  8. ^ "Google Doodle honors DNA researcher Har Gobind Khorana". usatoday.com. Retrieved 9 January 2018.
  9. ^ a b "Biochemist Har Gobind Khorana, whose UW work earned the Nobel Prize, dies". news.wisc.edu. Retrieved 2017-01-10.
  10. ^ Kresge, Nicole; Simoni, Robert D.; Hill, Robert L. (2009-05-29). "Total Synthesis of a Tyrosine Suppressor tRNA: the Work of H. Gobind Khorana". The Journal of Biological Chemistry. 284 (22): e5. ISSN 0021-9258. PMC 2685647.
  11. ^ "Gobind Khorana, MIT professor emeritus, dies at 89". MIT News. Retrieved 2018-01-09.
  12. ^ "MIT HG Khorana MIT laboratory".
  13. ^ a b c d e Zhao, Xiaojian; Ph.D, Edward J. W. Park (26 November 2013). "Asian Americans: An Encyclopedia of Social, Cultural, Economic, and Political History [3 volumes]: An Encyclopedia of Social, Cultural, Economic, and Political History". ABC-CLIO. Retrieved 9 January 2018 – via Google Books.
  14. ^ a b "H Gobind Khorana". 15 November 2011. Retrieved 9 January 2018 – via www.telegraph.co.uk.
  15. ^ "Har Gobind Khorana: Why Google honours him today". www.aljazeera.com. Retrieved 9 January 2018.
  16. ^ a b c d e "Gobind Khorana, MIT professor emeritus, dies at 89". mit.edu. Retrieved 9 January 2018.
  17. ^ a b "Gobind Khorana, MIT professor emeritus, dies at 89". mit.edu. Retrieved 9 January 2018.
  18. ^ a b "All you need to know about Har Gobind Ghorana, who Google is celebrating today with a Doodle". independent.co.uk. 8 January 2018. Retrieved 9 January 2018.
  19. ^ Kumar, Chandan. "H. Gobind Khorana - Biographical". Nobel Prize.org. Nobel Prize. Retrieved 9 January 2018.
  20. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  21. ^ https://books.google.ca/books?id=3AxIAgAAQBAJ&pg=PA653&dq=early+life+har+gobind+khorana&hl=en&sa=X&ved=0ahUKEwiNreuXt8vYAhVOxGMKHW7dB10Q6AEILDAB#v=onepage&q=early%20life%20har%20gobind%20khorana&f=false, p=653
  22. ^ RajBhandary, Uttam L. (14 December 2011). "Har Gobind Khorana (1922–2011)". Nature. Retrieved 9 January 2018.
  23. ^ a b RajBhandary, Uttam L. (14 December 2011). "Har Gobind Khorana (1922–2011)". Nature. Retrieved 9 January 2018.
  24. ^ https://biochem.ubc.ca/about/history/
  25. ^ Zhao, Xiaojian; Ph.D, Edward J. W. Park (26 November 2013). "Asian Americans: An Encyclopedia of Social, Cultural, Economic, and Political History [3 volumes]: An Encyclopedia of Social, Cultural, Economic, and Political History". ABC-CLIO. Retrieved 9 January 2018 – via Google Books.
  26. ^ Lindsten, Jan (9 January 1999). "Physiology Or Medicine, 1963-1970". World Scientific. Retrieved 9 January 2018 – via Google Books.
  27. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  28. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  29. ^ a b c "Har Gobind Khorana deciphered DNA and wrote the dictionary for our genetic language". vox.com. Retrieved 9 January 2018.
  30. ^ a b "Who Is Har Gobind Khorana? Why Google Is Celebrating the Nobel Prize-Winner". Time. Retrieved 9 January 2018.
  31. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  32. ^ a b "HG Khorana Nobel Lecture".
  33. ^ a b Khorana, H. G. (1979). "Total synthesis of a gene". Science. 203 (4381): 614–625. Bibcode:1979Sci...203..614K. doi:10.1126/science.366749. PMID 366749.
  34. ^ Kleppe, K.; Ohtsuka, E.; Kleppe, R.; Molineux, I.; Khorana, H. G. (1971). "Studies on polynucleotides *1, *2XCVI. Repair replication of short synthetic DNA's as catalyzed by DNA polymerases". Journal of Molecular Biology. 56 (2): 341–361. doi:10.1016/0022-2836(71)90469-4. PMID 4927950.
  35. ^ Wildenauer, D.; Khorana, H. G. (1977). "The preparation of lipid-depleted bacteriorhodopsin". Biochimica et Biophysica Acta. 466 (2): 315–324. doi:10.1016/0005-2736(77)90227-9. PMID 857886.
  36. ^ Ahuja, S.; Crocker, E.; Eilers, M.; Hornak, V.; Hirshfeld, A.; Ziliox, M.; Syrett, N.; Reeves, P. J.; Khorana, H. G.; Sheves, M.; Smith, S. O. (2009). "Location of the Retinal Chromophore in the Activated State of Rhodopsin*". Journal of Biological Chemistry. 284 (15): 10190–10201. doi:10.1074/jbc.M805725200. PMC 2665073. PMID 19176531.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  37. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  38. ^ H. Gobind Khorana - Biographical
  39. ^ "Fellowship of the Royal Society 1660-2015". London: Royal Society. Archived from the original on 15 July 2015. {{cite web}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help)
  40. ^ http://iusstf.org/story/53-50-Khorana-Program.html
  41. ^ Molina, Brett. "Google Doodle honors DNA researcher Har Gobind Khorana". USA Today. Retrieved January 9, 2018.
  42. ^ Musil, Steven (January 8, 2018). "Google Doodle honors biochemist Har Gobind Khorana". CBS Interactive Inc. Retrieved January 9, 2018.
  43. ^ https://www.washingtonpost.com/local/obituaries/h-gobind-khorana-biochemist-and-nobel-prize-winner-dies/2011/11/11/gIQAe4rIDN_story.html?utm_term=.d100f71144c1
  44. ^ https://www.washingtonpost.com/local/obituaries/h-gobind-khorana-biochemist-and-nobel-prize-winner-dies/2011/11/11/gIQAe4rIDN_story.html?utm_term=.d100f71144c1
  45. ^ https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html
  46. ^ https://www.washingtonpost.com/local/obituaries/h-gobind-khorana-biochemist-and-nobel-prize-winner-dies/2011/11/11/gIQAe4rIDN_story.html?utm_term=.d100f71144c1
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