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Har Gobind KhoranaForMemRS
Born	(1922-01-09)9 January 1922 Raipur, Multan, Punjab Province, British India
Died	9 November 2011(2011-11-09) (aged 89) Concord, Massachusetts, U.S.
Citizenship	United States
Education	Government College, Lahore Punjab University (BS, MS) University of Liverpool (PhD)
Known for	First to demonstrate the role of nucleotides in protein synthesis
Spouse	Esther Elizabeth Sibler
Awards	Nobel Prize in Medicine (1968) Gairdner Foundation International Award (1980) Louisa Gross Horwitz Prize Albert Lasker Award for Basic Medical Research Padma Vibhushan Willard Gibbs Award (1974)
Scientific career
Fields	Molecular biology
Institutions	MIT (1970–2007) University of Wisconsin–Madison (1960–70) University of British Columbia (1952–60) University of Cambridge (1950–52) Swiss Federal Institute of Technology, Zurich (1948–49)
Doctoral advisor	Roger J.S. Beer
Doctoral students	Shiladitya DasSarma
Signature

Har Gobind Khorana (9 January 1922 – 9 November 2011) was an Indian-American biochemist. 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.

Born in British India, Khorana served on the faculties of three universities in North America. He became a naturalized citizen of the United States in 1966, and received the National Medal of Science in 1987.

Biography

Har Gobind Khorana was born to Ganpatrai Khorana and Krishna Devi, in Raipur, a village in Multan, Punjab, British India, in a Punjabi Hindu Khatri family. The exact date of his birth is not certain but he believed that it might have been 9 January 1922; this date was later shown in some documents, and has been widely accepted. 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." The first four years of his education were provided under a tree, a spot that was, in effect, the only school in the village. He did not even own a pencil until age 6.

He attended D.A.V. (Dayanand Anglo-Vedic) High School in Multan and Government College, in Lahore. Later, he studied at the Punjab University in Lahore, with the assistance of scholarships, where he obtained a bachelor's degree in 1943 and a Master of Science degree in 1945.

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. The following year, he pursued postdoctoral studies with Professor Vladimir Prelog at ETH Zurich in Switzerland. He worked for nearly a year on alkaloid chemistry in an unpaid position.

His family moved to Delhi from Multan as refugees during the partition of India and Khorana was never to visit his place of birth after that. During a brief period in 1949, he was unable to find a job in Delhi. He returned to England on a fellowship to work with George Wallace Kenner and Alexander R. Todd on peptides and nucleotides. 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. Khorana was excited by the prospect of starting his own lab, a colleague later recalled. His mentor later said that the council had few facilities at the time but gave the researcher "all the freedom in the world". His work in British Columbia was on "nucleic acids and synthesis of many important biomolecules" according to the American Chemical Society.

In 1960 Khorana accepted a position as co-director of the University of Wisconsin–Madison's Institute for Enzyme Research He became a professor of biochemistry in 1962 and was named Conrad A. Elvehjem Professor of Life Sciences in 1964. 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". During his tenure at this university, he completed the work that led to sharing the Nobel Prize in 1968. 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."

He became a US citizen in 1966. Beginning in 1970, Khorana was the Alfred P. Sloan Professor of Biology and Chemistry at the Massachusetts Institute of Technology and later, a member of the Board of Scientific Governors at The Scripps Research Institute. He retired from MIT in 2007.

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

Ribonucleic acid (RNA) with two 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.

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

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. 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, as well as methods that anticipated the invention of polymerase chain reaction (PCR). 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. Later, his lab went on to study the structurally related visual pigment known as rhodopsin.

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."

Awards and honors

In addition to sharing the Nobel prize, Khorana was elected a member of the United States National Academy of Sciences in 1966, a member of the American Academy of Arts and Sciences in 1967, a member of the American Philosophical Society in 1973, and a Foreign Member of the Royal Society (ForMemRS) in 1978. The Indian government awarded Khorana the Padma Vibhushan in 1969.

In 2009, Khorana was hosted by the Khorana Program and honored at the 33rd Steenbock Symposium in Madison, Wisconsin.

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 American Academy of Achievement in 1971, 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.

On 9 January 2018, a Google Doodle celebrated the achievements of Har Gobind Khorana on what would have been his 96th birthday.

The "Khorana Program" by Government of India's Department of Biotechnology, the University of Wisconsin–Madison, Wisconsin–India Science and Technology Exchange Program (WINStep Forward/WSF) and the Indo-US Science and Technology Forum is a scholarship instituted in his name, since 2007 it has offered Indian students research opportunities at certain US universities. 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: 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 WINStep Forward adopted administration responsibilities for the Khorana program in 2007. 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.

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, but Khorana was survived by his other two children. 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."

References

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External links

• The Khorana Program
• 33rd Steenbock Symposium
• Remembering Har Gobind Khorana: University of Wisconsin Biochemistry Newsletter, adapted from article in Cell
• Har Gobind Khorana materials in the South Asian American Digital Archive (SAADA)
• Har Gobind Khorana (1922–2011): Pioneering Spirit (obituary)
• Har Gobind Khorana
• H. Gobind Khorana on Nobelprize.org
• HG Khorana Birth Centenary Lecture (Jan 9, 2022)

v t e Laureates of the Nobel Prize in Physiology or Medicine
1901–1925	1901: Emil Behring 1902: Ronald Ross 1903: Niels Finsen 1904: Ivan Pavlov 1905: Robert Koch 1906: Camillo Golgi / Santiago Ramón y Cajal 1907: Alphonse Laveran 1908: Élie Metchnikoff / Paul Ehrlich 1909: Emil Kocher 1910: Albrecht Kossel 1911: Allvar Gullstrand 1912: Alexis Carrel 1913: Charles Richet 1914: Róbert Bárány 1915 1916 1917 1918 1919: Jules Bordet 1920: August Krogh 1921 1922: Archibald Hill / Otto Meyerhof 1923: Frederick Banting / John Macleod 1924: Willem Einthoven 1925
1926–1950	1926: Johannes Fibiger 1927: Julius Wagner-Jauregg 1928: Charles Nicolle 1929: Christiaan Eijkman / Frederick Gowland Hopkins 1930: Karl Landsteiner 1931: Otto Warburg 1932: Charles Scott Sherrington / Edgar Adrian 1933: Thomas Morgan 1934: George Whipple / George Minot / William Murphy 1935: Hans Spemann 1936: Henry Dale / Otto Loewi 1937: Albert Szent-Györgyi 1938: Corneille Heymans 1939: Gerhard Domagk 1940 1941 1942 1943: Henrik Dam / Edward Doisy 1944: Joseph Erlanger / Herbert Gasser 1945: Alexander Fleming / Ernst Chain / Howard Florey 1946: Hermann Muller 1947: Carl Cori / Gerty Cori / Bernardo Houssay 1948: Paul Müller 1949: Walter Hess / António Egas Moniz 1950: Edward Kendall / Tadeusz Reichstein / Philip Hench
1951–1975	1951: Max Theiler 1952: Selman Waksman 1953: Hans Krebs / Fritz Lipmann 1954: John Enders / Thomas Weller / Frederick Robbins 1955: Hugo Theorell 1956: André Cournand / Werner Forssmann / Dickinson W. Richards 1957: Daniel Bovet 1958: George Beadle / Edward Tatum / Joshua Lederberg 1959: Severo Ochoa / Arthur Kornberg 1960: Frank Burnet / Peter Medawar 1961: Georg von Békésy 1962: Francis Crick / James Watson / Maurice Wilkins 1963: John Eccles / Alan Hodgkin / Andrew Huxley 1964: Konrad Bloch / Feodor Lynen 1965: François Jacob / André Lwoff / Jacques Monod 1966: Francis Rous / Charles B. Huggins 1967: Ragnar Granit / Haldan Hartline / George Wald 1968: Robert W. Holley / Har Khorana / Marshall Nirenberg 1969: Max Delbrück / Alfred Hershey / Salvador Luria 1970: Bernard Katz / Ulf von Euler / Julius Axelrod 1971: Earl Sutherland Jr. 1972: Gerald Edelman / Rodney Porter 1973: Karl von Frisch / Konrad Lorenz / Nikolaas Tinbergen 1974: Albert Claude / Christian de Duve / George Palade 1975: David Baltimore / Renato Dulbecco / Howard Temin
1976–2000	1976: Baruch Blumberg / Daniel Gajdusek 1977: Roger Guillemin / Andrew Schally / Rosalyn Yalow 1978: Werner Arber / Daniel Nathans / Hamilton O. Smith 1979: Allan Cormack / Godfrey Hounsfield 1980: Baruj Benacerraf / Jean Dausset / George Snell 1981: Roger Sperry / David H. Hubel / Torsten Wiesel 1982: Sune Bergström / Bengt I. Samuelsson / John Vane 1983: Barbara McClintock 1984: Niels Jerne / Georges Köhler / César Milstein 1985: Michael Brown / Joseph L. Goldstein 1986: Stanley Cohen / Rita Levi-Montalcini 1987: Susumu Tonegawa 1988: James W. Black / Gertrude B. Elion / George H. Hitchings 1989: J. Michael Bishop / Harold E. Varmus 1990: Joseph Murray / E. Donnall Thomas 1991: Erwin Neher / Bert Sakmann 1992: Edmond Fischer / Edwin G. Krebs 1993: Richard J. Roberts / Phillip Sharp 1994: Alfred G. Gilman / Martin Rodbell 1995: Edward B. Lewis / Christiane Nüsslein-Volhard / Eric F. Wieschaus 1996: Peter C. Doherty / Rolf M. Zinkernagel 1997: Stanley B. Prusiner 1998: Robert F. Furchgott / Louis Ignarro / Ferid Murad 1999: Günter Blobel 2000: Arvid Carlsson / Paul Greengard / Eric Kandel
2001–present	2001: Leland H. Hartwell / Tim Hunt / Paul Nurse 2002: Sydney Brenner / H. Robert Horvitz / John E. Sulston 2003: Paul Lauterbur / Peter Mansfield 2004: Richard Axel / Linda B. Buck 2005: Barry Marshall / Robin Warren 2006: Andrew Fire / Craig Mello 2007: Mario Capecchi / Martin Evans / Oliver Smithies 2008: Harald zur Hausen / Luc Montagnier / Françoise Barré-Sinoussi 2009: Elizabeth Blackburn / Carol W. Greider / Jack W. Szostak 2010: Robert G. Edwards 2011: Bruce Beutler / Jules A. Hoffmann / Ralph M. Steinman (posthumously) 2012: John Gurdon / Shinya Yamanaka 2013: James Rothman / Randy Schekman / Thomas C. Südhof 2014: John O'Keefe / May-Britt Moser / Edvard Moser 2015: William C. Campbell / Satoshi Ōmura / Tu Youyou 2016: Yoshinori Ohsumi 2017: Jeffrey C. Hall / Michael Rosbash / Michael W. Young 2018: James P. Allison / Tasuku Honjo 2019: Gregg L. Semenza / Peter J. Ratcliffe / William Kaelin Jr. 2020: Harvey J. Alter / Michael Houghton / Charles M. Rice 2021: David Julius / Ardem Patapoutian 2022: Svante Pääbo 2023: Katalin Karikó / Drew Weissman 2024: Victor Ambros / Gary Ruvkun

• 1922 births
• 2011 deaths
• University of the Punjab alumni
• Members of the United States National Academy of Sciences
• Alumni of the University of Liverpool
• Nobel laureates in Physiology or Medicine
• Members of the Pontifical Academy of Sciences
• National Medal of Science laureates
• Foreign members of the Royal Society
• Foreign members of the Russian Academy of Sciences
• Massachusetts Institute of Technology School of Science faculty
• Scripps Research
• Recipients of the Padma Vibhushan in science & engineering
• Academic staff of the University of British Columbia
• Recipients of the Albert Lasker Award for Basic Medical Research
• People from Khanewal District
• Government College University, Lahore alumni
• Naturalized citizens of the United States
• People who lost Indian citizenship
• Indian emigrants to the United States
• American people of Punjabi descent
• American Nobel laureates
• Members of the American Philosophical Society
• Dayanand Anglo-Vedic Schools System alumni
• People from Punjab Province (British India)
• Academic staff of ETH Zurich