ImportMaster at 19:17, 9 May 2022

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{{Short description|Pakistani-American molecular biologist}}
{{EngvarB|date=January 2018}}
{{Use dmy dates|date=August 2016}}
{{Infobox scientist
| name = Har Gobind Khorana
| image = Har Gobind Khorana nobel.jpg
| caption =
| birth_date = {{birth date |df=yes|1922|1|09}}
| birth_place = [[Raipur, Pakistan|Raipur]], [[Multan]], [[Punjab Province (British India)|Punjab Province]], [[British India]] (present day [[Punjab, Pakistan|Punjab]], [[Pakistan]])
| death_date = {{death date and age |df=yes|2011|11|09 |1922|01|09}}
| death_place = [[Concord, Massachusetts]], U.S.
| citizenship = [[United States]]
| spouse = Esther Elizabeth Sibler
| field = [[Molecular biology]]
| work_institution = {{Plainlist|
* [[MIT]] (1970–2007)
* [[University of Wisconsin–Madison]] (1960–70)
* [[University of British Columbia]] (1952–60)
* [[University of Cambridge]] (1950–52)
* [[ETH Zurich|Swiss Federal Institute of Technology, Zurich]] (1948–49)}}
| alma_mater = {{plainlist|
* [[Government College University, Lahore]]
* [[University of the Punjab|Punjab University]] ([[B.S.|BS]], [[M.S.|MS]])
* [[University of Liverpool]] ([[PhD]]) }}
| doctoral_advisor = Roger J.S. Beer
| doctoral_students = [[Shiladitya DasSarma]]
| known_for = First to demonstrate the role of [[nucleotides]] in [[protein synthesis]]
| prizes = {{Plainlist|
* [[Nobel Prize in Medicine]] (1968)
* [[Gairdner Foundation International Award]] (1980)
* [[Louisa Gross Horwitz Prize]]
* [[Fellow of the Royal Society|ForMemRS]] (1978)
* [[Albert Lasker Award for Basic Medical Research]]
* [[Padma Vibhushan]]
* [[Willard Gibbs Award]] (1974)}}
| signature = Har Gobind Khorana signature.jpg
| signature_alt = Har Gobind Khorana signature
}}

'''Har Gobind Khorana''' (9 January 1922 – 9 November 2011)<ref>{{cite journal |last1=Caruthers |first1=M. |last2=Wells |first2=R. |doi=10.1126/science.1217138 |title=Har Gobind Khorana (1922-2011) |journal=Science |volume=334 |issue=6062 |pages=1511 |year=2011 |pmid=22174242|bibcode=2011Sci...334.1511C |s2cid=206539004 }}</ref><ref>{{cite journal |last1=Rajbhandary |first1=U. L. |title=Har Gobind Khorana (1922–2011) |doi=10.1038/480322a |journal=Nature |volume=480 |issue=7377 |pages=322 |year=2011 |pmid=22170673|bibcode=2011Natur.480..322R|doi-access=free }}</ref><ref>{{Cite news|url=https://www.nytimes.com/2011/11/14/us/h-gobind-khorana-1968-nobel-winner-for-rna-research-dies.html|title=H. Gobind Khorana, 1968 Nobel Winner biochemist for RNA Research, Dies|last=Gellene|first=Denise|date=2011-11-14|work=The New York Times|access-date=2018-01-09|language=en-US|issn=0362-4331}}</ref> was an [[Indian Americans|Indian-American]] biochemist.<ref name=NobelBio/> While on the faculty of the [[University of Wisconsin–Madison]], he shared the 1968 [[Nobel Prize for Physiology or Medicine]] with [[Marshall W. Nirenberg]] and [[Robert W. Holley]] for research that showed the order of [[nucleotides]] in [[nucleic acids]], which carry the [[genetic code]] of the cell and 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|date=14 June 2018}}</ref><ref name="PLoSBio">{{Cite journal|title=Har Gobind Khorana (1922–2011): Pioneering Spirit|last=Sakmar|first=Thomas P.|date=2012-12-02|journal=PLOS Biology|volume=10|issue=2|pages=e1001273|language=en-US|issn=1545-7885|doi=10.1371/journal.pbio.1001273 |pmc=3283548|doi-access=free}}</ref>

Born in [[British India]], modern-day [[Pakistan]], Khorana served on the faculties of three universities in [[North America]]. He became a [[naturalized citizen]] of the [[United States]] in 1966,<ref name="brit">{{cite encyclopedia |url=http://www.britannica.com/EBchecked/topic/316846/Har-Gobind-Khorana |title=Har Gobind Khorana: American biochemist|encyclopedia=Encyclopaedia Britannica}}</ref> and 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|access-date=9 January 2018}}</ref>

==Biography==
Khorana was born to Krishna Devi Khorana and Ganpat Rai Khorana, in [[Raipur, Pakistan|Raipur]], a village in [[Multan]], [[Punjab region|Punjab]], British India in a [[Punjabis|Punjabi]] [[Hindu]] family.<ref name="auto">{{cite book|url=https://books.google.com/books?id=3AxIAgAAQBAJ&q=early+life+har+gobind+khorana&pg=PA653|title=Asian Americans: An Encyclopedia of Social, Cultural, Economic, and Political History [3 volumes]: An Encyclopedia of Social, Cultural, Economic, and Political History|first1=Xiaojian|last1=Zhao|first2=Edward J. W. Park|last2=Ph.D|date=26 November 2013|publisher=ABC-CLIO|access-date=9 January 2018|via=Google Books|isbn=9781598842401}}</ref> The exact date of his birth is not certain but he believed that it might have been 9 January 1922;<ref name="telegraph.co.uk">{{cite news|url=https://www.telegraph.co.uk/news/obituaries/science-obituaries/8892230/H-Gobind-Khorana.html |archive-url=https://ghostarchive.org/archive/20220112/https://www.telegraph.co.uk/news/obituaries/science-obituaries/8892230/H-Gobind-Khorana.html |archive-date=12 January 2022 |url-access=subscription |url-status=live|title=H Gobind Khorana|date=15 November 2011|access-date=7 January 2019|work=The Telegraph}}{{cbignore}}</ref> this date was later shown in some documents, and has been widely accepted.<ref>{{cite web|url=http://www.aljazeera.com/news/2018/01/har-gobind-khorana-google-honours-today-180108134719847.html|title=Har Gobind Khorana: Why Google honours him today|publisher=Al Jazeera|access-date=9 January 2018}}</ref> He was the youngest of five children. His father was a [[patwari]], a village agricultural taxation clerk in the British 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."<ref name="mit.edu">{{cite web|url=https://news.mit.edu/2011/obit-khorana-1110|title=Gobind Khorana, MIT professor emeritus, dies at 89|website=mit.edu|access-date=9 January 2018}}</ref> The first four years of his education were provided under a tree, a spot that was, in effect, the only school in the village.<ref name="auto" /> He did not even own a pencil until age 6.<ref name="Sarkar" />

He attended D.A.V. ([[Dayanand Anglo Vedic|Dayanand Anglo-Vedic]]) High School in [[Multan]], in [[Punjab, Pakistan|West Punjab]].<ref name="auto" /> Later, he studied at the [[University of the Punjab|Punjab University]] in [[Lahore]], with the assistance of scholarships, where he obtained a bachelor's degree in 1943<ref name="mit.edu" /> and a Master of Science degree in 1945.<ref name=NobelBio>{{cite web|last1=Kumar|first1=Chandan|url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/1968/khorana-bio.html|title=H. Gobind Khorana – Biographical|website=Nobel Prize.org|publisher=Nobel Prize|access-date=9 January 2018}}</ref><ref name="independent.co.uk">{{cite web|url=https://www.independent.co.uk/news/world/americas/har-gobind-ghorana-birthday-who-is-he-google-doodle-dna-science-life-career-a8148666.html |archive-url=https://ghostarchive.org/archive/20220509/https://www.independent.co.uk/news/world/americas/har-gobind-ghorana-birthday-who-is-he-google-doodle-dna-science-life-career-a8148666.html |archive-date=9 May 2022 |url-access=subscription |url-status=live|title=All you need to know about Har Gobind Ghorana, who Google is celebrating today with a Doodle|date=8 January 2018|website=Independent|access-date=9 January 2018}}</ref>

Khorana lived in British India until 1945, when he moved to England to study organic chemistry at the [[University of Liverpool]] on a Government of India Fellowship. He received his PhD in 1948 advised by Roger J. S. Beer.<ref name=CenAcs>{{cite web|url=https://cen.acs.org/articles/89/i47/Har-Gobind-Khorana-Dies-89.html|title=Har Gobind Khorana Dies At 89 – November 21, 2011 Issue – Vol. 89 Issue 47 – Chemical & Engineering News|first=Susan J.|last=Ainsworth|website=cen.acs.org|access-date=9 January 2018}}</ref><ref name="Zhao 2014 pp. 652-655">{{cite book |first=Benjamin C. |last=Zulueta |editor-last=Zhao |editor-first=Xiaojian |title=Asian Americans: An Encyclopedia of Social, Cultural, Economic, and Political History |chapter=Khorana, Har Gobind (1922–2011) |publisher=Greenwood, an imprint of ABC-CLIO, LLC |location=Santa Barbara, California |year=2013 |chapter-url=https://books.google.com/books?id=3AxIAgAAQBAJ&pg=PA653 |isbn=9781598842401 |pages=652–655 |oclc=911399074 }}</ref><ref name="auto1"/><ref name="mit.edu"/> The following year, he pursued postdoctoral studies with Professor [[Vladimir Prelog]] at [[ETH Zurich]] in [[Switzerland]].<ref name="mit.edu"/> He worked for nearly a year on alkaloid chemistry in an unpaid position.<ref name="auto"/><ref name="auto1">{{cite journal|title=Har Gobind Khorana (1922–2011)|first=Uttam L.|last=RajBhandary|date=14 December 2011|journal=Nature|volume=480|issue=7377|pages=322|pmid=22170673|doi=10.1038/480322a|bibcode=2011Natur.480..322R|doi-access=free}}</ref>

During a brief period in 1949, he was unable to find a job in his original home area in the Punjab.<ref name="auto"/> He returned to England on a fellowship to work with [[George Wallace Kenner]] and [[Alexander R. Todd]] on peptides and nucleotides.<ref name="auto1"/> He stayed in Cambridge from 1950 until 1952.

He moved to Vancouver, [[British Columbia]], with his family in 1952 after accepting a position with the British Columbia Research Council at [[University of British Columbia]].<ref name="mit.edu"/><ref>{{cite web|url=https://biochem.ubc.ca/about/history/|title=History – Department of Biochemistry|website=biochem.ubc.ca|access-date=9 January 2018}}</ref> Khorana was excited by the prospect of starting his own lab, a colleague later recalled.<ref name="auto"/> His mentor later said that the Council had few facilities at the time but gave the researcher "all the freedom in the world".<ref>{{cite book|url=https://books.google.com/books?id=O5jQ-GRTvPUC&q=mahan+singh+professor+researcher+Punjab+university&pg=PA370|title=Physiology Or Medicine, 1963–1970|first=Jan|last=Lindsten|date=9 January 1999|publisher=World Scientific|access-date=9 January 2018|via=Google Books|isbn=9789810234126}}</ref> His work in British Columbia was on "nucleic acids and synthesis of many important biomolecules" according to the [[American Chemical Society]].<ref name=CenAcs/>

In 1960 Khorana accepted a position as co-director of the Institute for Enzyme research at the Institute for Enzyme Research at the [[University of Wisconsin]] at Madison.<ref name=CenAcs/><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> He became a professor of biochemistry in 1962 and was named [[Conrad Elvehjem|Conrad A. Elvehjem]] Professor of Life Sciences at 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 |doi=10.1016/S0021-9258(19)82144-0 |issn=0021-9258 |pmc=2685647|doi-access=free }}</ref> 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.<ref name=CenAcs /> During his tenure at this University, he completed the work that led to sharing the Nobel prize. The Nobel web site states that it was "for their interpretation of the genetic code and its function in protein synthesis". Har Gobind Khorana's role is stated as follows: he "made important contributions to this field by building different RNA chains with the help of enzymes. Using these enzymes, he was able to produce proteins. The amino acid sequences of these proteins then solved the rest of the puzzle."<ref>{{cite web|url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/1968/khorana-facts.html|title=H. Gobind Khorana – Facts|website=www.nobelprize.org|access-date=9 January 2018}}</ref>

He became a US citizen in 1966.<ref name="vox.com" /> Beginning in 1970, Khorana was the [[Alfred P. Sloan]] Professor of Biology and Chemistry at the [[Massachusetts Institute of Technology]]<ref name="time.com" /><ref name="mit.edu" /><ref>{{cite web |url=http://web.mit.edu/chemistry/www/faculty/khorana.html |title=MIT HG Khorana MIT laboratory}}</ref> and later, a member of the Board of Scientific Governors at [[The Scripps Research Institute]]. He retired from MIT in 2007.<ref name="vox.com" />

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

==Research==
[[RNA|Ribonucleic acid]] (RNA) with two repeating units (UCUCUCU → UCU CUC UCU) produced two alternating [[amino acid]]s. 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 [[dipeptide]]s and [[tripeptide]]s thus revealing that UAG, UAA, and UGA are [[stop codon]]s.<ref name="nobel"/>

Their [[Nobel Prize|Nobel]] lecture was delivered on 12 December 1968.<ref name="nobel">{{cite web |url=http://nobelprize.org/nobel_prizes/medicine/laureates/1968/khorana-lecture.html |title=HG Khorana Nobel Lecture}}</ref> Khorana was the first scientist to chemically synthesize [[oligonucleotide]]s.<ref name="doi10.1126/science.366749">{{cite journal |last1=Khorana |first1=H. G. |author-link=Har Gobind Khorana |title=Total synthesis of a gene |doi=10.1126/science.366749 |journal=Science |volume=203 |issue=4381 |pages=614–625 |year=1979 |pmid=366749 |bibcode=1979Sci...203..614K |url=http://dspace.mit.edu/bitstream/1721.1/104342/1/12045_2012_Article_134.pdf }}</ref> This achievement, in the 1970s, was also the world's first synthetic gene; in later years, the process has become widespread.<ref name="time.com">{{cite magazine|url=http://time.com/5094695/google-doodle-har-gobind-khorana/|title=Who Is Har Gobind Khorana? Why Google Is Celebrating the Nobel Prize-Winner|magazine=Time|access-date=9 January 2018}}</ref> Subsequent scientists referred to his research while advancing genome editing with the CRISPR/Cas9 system.<ref name="vox.com">{{cite web|url=https://www.vox.com/2018/1/9/16862980/google-doodle-har-gobind-khorana-genetics-dna|title=Har Gobind Khorana deciphered DNA and wrote the dictionary for our genetic language|website=vox.com|access-date=9 January 2018|date=9 January 2018}}</ref>

===Subsequent research===
After years of work, he was the first in the world to complete the total synthesis of a functional gene outside a living organism in 1972.<ref name="Sarkar">{{cite news |last1=Sarkar |first1=Sahotra |title=Har Gobind Khorana: The chemist who cracked DNA’s code and made the first artificial gene was born into poverty 100 years ago in an Indian village |url=https://theconversation.com/har-gobind-khorana-the-chemist-who-cracked-dnas-code-and-made-the-first-artificial-gene-was-born-into-poverty-100-years-ago-in-an-indian-village-178390 |access-date=7 April 2022 |work=The Conversation |date=5 April 2022}}</ref> He did this by extending 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,<ref name="doi10.1126/science.366749"/> as well as methods that anticipated the invention of [[polymerase chain reaction]] (PCR).<ref>{{cite journal |last1=Kleppe |first1=K. |last2=Ohtsuka |first2=E. |last3=Kleppe |first3=R. |last4=Molineux |first4=I. |last5=Khorana |first5=H. G. |author-link5=Har Gobind Khorana |title=Studies on polynucleotides *1, *2XCVI. Repair replication of short synthetic DNA's as catalyzed by DNA polymerases |doi=10.1016/0022-2836(71)90469-4 |journal=Journal of Molecular Biology |volume=56 |issue=2 |pages=341–361 |year=1971 |pmid=4927950 }}</ref> 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 synthesis|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.<ref>{{cite journal|last1=Wildenauer |first1=D. |last2=Khorana |first2=H. G. |author-link2=Har Gobind Khorana |title=The preparation of lipid-depleted bacteriorhodopsin |journal=Biochimica et Biophysica Acta (BBA) - Biomembranes |volume=466 |issue=2 |pages=315–324 |year=1977 |pmid=857886 |doi=10.1016/0005-2736(77)90227-9}}</ref> Later, his lab went on to study the structurally related visual pigment known as [[rhodopsin]].<ref>{{cite journal |last1=Ahuja |first1=S. |last2=Crocker |first2=E. |last3=Eilers |first3=M. |last4=Hornak |first4=V. |last5=Hirshfeld |first5=A. |last6=Ziliox |first6=M. |last7=Syrett |first7=N. |last8=Reeves |first8=P. J. |last9=Khorana |first9=H. G. |author-link9=Har Gobind Khorana |last10=Sheves |doi=10.1074/jbc.M805725200 |first10=M. |last11=Smith |first11=S. O. |title=Location of the Retinal Chromophore in the Activated State of Rhodopsin* |journal=Journal of Biological Chemistry |volume=284 |issue=15 |pages=10190–10201 |year=2009 |pmid=19176531 |pmc=2665073|doi-access=free }}</ref>

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."<ref name=CenAcs/><ref name=NobelBio/>

==Awards and honors==
[[File:Har Gobind Khorana receives NIH lecture award.jpg|thumb|Har Gobind Khorana receiving NIH lecture award.]]
In addition to sharing the [[Nobel prize]],<ref name="independent.co.uk"/> Khorana was elected as [[List of Fellows of the Royal Society elected in 1978|Foreign Member of the Royal Society (ForMemRS) in 1978]].<ref name="fellows">{{cite web |title=Fellowship of the Royal Society 1660–2015 |url=https://docs.google.com/spreadsheets/d/1RVVZY00MZNrK2YCTTzVrbTFH2t3RxoAZah128gQR-NM/pubhtml |publisher=[[Royal Society]] |location=London |archive-url=https://web.archive.org/web/20151015185820/https://docs.google.com/spreadsheets/d/1RVVZY00MZNrK2YCTTzVrbTFH2t3RxoAZah128gQR-NM/pubhtml |archive-date=15 October 2015 |url-status=dead |df=dmy-all}}</ref> In 2007, the University of Wisconsin–Madison, the Government of India (DBT [[Department of Biotechnology]]), and the Indo-US Science and Technology Forum jointly created the Khorana Program. 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:<ref>{{cite web |url=http://iusstf.org/story/53-50-Khorana-Program.html |title=Khorana Program for Scholars |website=iusstf.org |publisher=Indo-US Science and Technology Forum |location=New Delhi, India |access-date=2018-01-09}}</ref> 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. The Wisconsin–India Science and Technology Exchange Program (WINStep Forward, WSF) adopted administration responsibilities for the Khorana program in 2007.<ref>{{cite web |url=https://www.winstepforward.org/history/ |title=History |website=winstepforward.org |publisher=WINStep Forward |location=Madison, WI |access-date=2018-06-26}}</ref> WINStep Forward was jointly created by Drs. Aseem Ansari and Ken Shapiro at the University of Wisconsin–Madison. WINStep Forward also administers the nationally competitive S.N. Bose Programs for Indian and American students, respectively, to promote both fundamental and applied research not only in biotechnology but broadly across all STEM (science, technology, engineering, and mathematics) fields, including medicine, pharmacy, agriculture, wildlife and climate change.

In 2009, Khorana was hosted by the Khorana Program and honored at the 33rd Steenbock Symposium in [[Madison, Wisconsin]].<ref name="news.wisc.edu"/>

Other honors included the Louisa Gross Horwitz Prize from [[Columbia University]] and the Lasker Foundation Award for Basic Medical Research, both in 1969, the Golden Plate Award of the [[Academy of Achievement|American Academy of Achievement]] in 1971,<ref>{{cite web|title= Golden Plate Awardees of the American Academy of Achievement |website=www.achievement.org|publisher=[[American Academy of Achievement]]|url=https://achievement.org/our-history/golden-plate-awards/#science-exploration}}</ref> 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.<ref name="mit.edu"/>

On 9 January 2018, a [[Google Doodle]] celebrated the achievements<ref>{{cite web|url=https://www.google.com/doodles/har-gobind-khoranas-96th-birthday|title=Har Gobind Khorana's 96th Birthday|website=www.google.com}}</ref> of Har Gobind Khorana on what would have been his 96th birthday.<ref name="Google 2018">{{cite web | title=Har Gobind Khorana's 96th Birthday | website=Google | date=2018-01-09 | url=https://www.google.com/doodles/har-gobind-khoranas-96th-birthday | access-date=2018-01-10}}</ref><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 |first=Brett |last=Molina |publisher=USA Today|access-date=9 January 2018}}</ref>

==Death==
Khorana died on 9 November 2011, in [[Concord, Massachusetts]], at the age of 89. His wife, Esther, and daughter, Emily Anne, had died earlier,<ref name=CenAcs/> but Khorana was survived by his other two children.<ref name="telegraph.co.uk"/> Julia Elizabeth later wrote about her father's work as a professor: "Even while doing all this research, he was always really interested in education, in students and young people."<ref name="mit.edu"/>

==References==
{{Reflist|30em}}

==External links==
* [https://web.archive.org/web/20170322214206/https://biochem.wisc.edu/khorana_program The Khorana Program]
* [https://biochem.wisc.edu/symposia/steenbock/33rd 33rd Steenbock Symposium]
* [https://biochem.wisc.edu/sites/default/files/About_Us/newsletters/2012BiochemNewsletter.pdf Remembering Har Gobind Khorana: University of Wisconsin Biochemistry Newsletter, adapted from article in ''Cell'']
* [http://www.saadigitalarchive.org/search/har%20gobind%20khorana Har Gobind Khorana materials in the South Asian American Digital Archive (SAADA)]
* [http://www.plosbiology.org/article/fetchObjectAttachment.action?uri=info%3Adoi%2F10.1371%2Fjournal.pbio.1001273&representation=PDF Har Gobind Khorana (1922–2011): Pioneering Spirit (obituary)]
*[http://www.universetale.com/har-gobind-khorana-the-nobel-laureate/ Har Gobind Khorana]
* {{Nobelprize|name=H. Gobind Khorana}}

{{Nobel Prize in Physiology or Medicine Laureates 1951-1975}}
{{1968 Nobel Prize winners}}
{{Winners of the National Medal of Science|biological}}
{{Padma Vibhushan Awards}}

{{Authority control}}
{{DEFAULTSORT:Khorana, Har Gobind}}
[[Category:1922 births]]
[[Category:2011 deaths]]
[[Category:University of the Punjab alumni]]
[[Category:Members of the United States National Academy of Sciences]]
[[Category:Alumni of the University of Liverpool]]
[[Category:Nobel laureates in Physiology or Medicine]]
[[Category:Punjabi academics]]
[[Category:Members of the Pontifical Academy of Sciences]]
[[Category:National Medal of Science laureates]]
[[Category:Foreign Members of the Royal Society]]
[[Category:Foreign Members of the Russian Academy of Sciences]]
[[Category:Massachusetts Institute of Technology School of Science faculty]]
[[Category:Scripps Research]]
[[Category:Recipients of the Padma Vibhushan in science & engineering]]
[[Category:University of British Columbia faculty]]
[[Category:Recipients of the Albert Lasker Award for Basic Medical Research]]
[[Category:People from Khanewal District]]
[[Category:Government College University, Lahore alumni]]
[[Category:People with acquired American citizenship]]
[[Category:American people of Punjabi descent]]
[[Category:American Nobel laureates]]
[[Category:Indian emigrants to the United States]]

Har Gobind Khorana - Revision history

ImportMaster at 19:17, 9 May 2022