Har Gobind Khorana

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Har Gobind Khorana
Har Gobind Khorana nobel.jpg
Born(1922-01-09)9 January 1922
Died9 November 2011(2011-11-09) (aged 89)
Residence British India
United States
Citizenship United States
Alma mater
Known forFirst to demonstrate the role of nucleotides in protein synthesis
Scientific career
Fields Molecular biology
Doctoral advisor Roger J.S. Beer
Doctoral students Shiladitya DasSarma
Har Gobind Khorana signature.jpg

Har Gobind Khorana (9 January 1922 – 9 November 2011) [1] [2] [3] was an Indian-American biochemist. [4] 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. [5] [6]

University of Wisconsin–Madison Public university in Wisconsin, USA

The University of Wisconsin–Madison is a public research university in Madison, Wisconsin. Founded when Wisconsin achieved statehood in 1848, UW–Madison is the official state university of Wisconsin, and the flagship campus of the University of Wisconsin System. It was the first public university established in Wisconsin and remains the oldest and largest public university in the state. It became a land-grant institution in 1866. The 933-acre (378 ha) main campus, located on the shores of Lake Mendota, includes four National Historic Landmarks. The University also owns and operates a historic 1,200-acre (486 ha) arboretum established in 1932, located 4 miles (6.4 km) south of the main campus.

Robert W. Holley American biochemist

Robert William Holley was an American biochemist. He shared the Nobel Prize in Physiology or Medicine in 1968 for describing the structure of alanine transfer RNA, linking DNA and protein synthesis.

Genetic code scheme, defines how sequences of codons specify which amino acid will be added next during protein synthesis; set of rules by which information encoded within genetic material (DNA or mRNA sequences) is translated into proteins by living cells

The genetic code is the set of rules used by living cells to translate information encoded within genetic material into proteins. Translation is accomplished by the ribosome, which links amino acids in an order specified by messenger RNA (mRNA), using transfer RNA (tRNA) molecules to carry amino acids and to read the mRNA three nucleotides at a time. The genetic code is highly similar among all organisms and can be expressed in a simple table with 64 entries.


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, [7] and received the National Medal of Science in 1987. [8]

United States Federal republic in North America

The United States of America (USA), commonly known as the United States or America, is a country comprising 50 states, a federal district, five major self-governing territories, and various possessions. At 3.8 million square miles, the United States is the world's third or fourth largest country by total area and is slightly smaller than the entire continent of Europe's 3.9 million square miles. With a population of over 327 million people, the U.S. is the third most populous country. The capital is Washington, D.C., and the largest city by population is New York City. Forty-eight states and the capital's federal district are contiguous in North America between Canada and Mexico. The State of Alaska is in the northwest corner of North America, bordered by Canada to the east and across the Bering Strait from Russia to the west. The State of Hawaii is an archipelago in the mid-Pacific Ocean. The U.S. territories are scattered about the Pacific Ocean and the Caribbean Sea, stretching across nine official time zones. The extremely diverse geography, climate, and wildlife of the United States make it one of the world's 17 megadiverse countries.

National Medal of Science award

The National Medal of Science is an honor bestowed by the President of the United States to individuals in science and engineering who have made important contributions to the advancement of knowledge in the fields of behavioral and social sciences, biology, chemistry, engineering, mathematics and physics. The twelve member presidential Committee on the National Medal of Science is responsible for selecting award recipients and is administered by the National Science Foundation (NSF).


Khorana was born to Krishna Devi Khorana and Ganpat Rai Khorana, in Raipur, a village in Punjab, British India (now in present-day Pakistan) in a Hindu family. [9] [10] The exact date of his birth is not certain but he believed that it might have been 9 January 1922; [11] this date was later shown in some documents, and has been widely accepted. [12] 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." [13] The first four years of his education were provided under a tree, a spot that was, in effect, the only school in the village. [10]

Raipur, Pakistan village in Punjab, Pakistan

Raipur, is a village in the Sowawa Tehsil of Jhelum District in Punjab, Pakistan. It is famous as the birthplace of Nobel Laureate Har Gobind Khorana.

Pakistan federal parliamentary constitutional republic in South Asia

Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the world’s sixth-most populous country with a population exceeding 212,742,631 people. In area, it is the 33rd-largest country, spanning 881,913 square kilometres. Pakistan has a 1,046-kilometre (650-mile) coastline along the Arabian Sea and Gulf of Oman in the south and is bordered by India to the east, Afghanistan to the west, Iran to the southwest, and China in the far northeast. It is separated narrowly from Tajikistan by Afghanistan's Wakhan Corridor in the northwest, and also shares a maritime border with Oman.

He attended D.A.V. High School in Multan, in West Punjab. [10] Later, he studied at the Punjab University in Lahore, with the assistance of scholarships, where he obtained a bachelor's degree in 1943 [13] and a Master of Science degree in 1945. [4] [14]

Multan City in Punjab

Multan is a city in Punjab, Pakistan. Located on the banks of the Chenab River, Multan is Pakistan's 7th largest city, and is the major cultural and economic centre of southern Punjab.

Punjab, Pakistan Province in Pakistan

Punjab is Pakistan's second largest province by area, after Balochistan, and it is the most populated province, with an estimated population of 110,012,442 as of 2017. Forming the bulk of the transnational Punjab region, it is bordered by the Pakistan provinces of Sindh, Balochistan, and Khyber Pakhtunkhwa, the enclave of Islamabad, and Azad Kashmir. It also shares borders with the Indian states of Punjab, Rajasthan and Jammu and Kashmir. The provincial capital of Punjab is the city Lahore, a cultural, historical, economic and cosmopolitan centre of Pakistan where the country's cinema industry, and much of its fashion industry, are based.

University of the Punjab public sector university primarily located in Lahore, Punjab, Pakistan

The University of the Punjab, also referred to as Punjab University, is a public research university located in Lahore, Punjab, Pakistan. Punjab University is the oldest public university in Pakistan. With multiple campuses in Gujranwala, Jhelum, and Khanspur, the university was formally established by the British Government after convening the first meeting for establishing higher education institutions in October 1882 at Simla. Punjab University was the fourth university to be established by the British colonial authorities in the Indian subcontinent; the first three universities were established in other parts of British India.

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. [15] [16] [17] [13] The following year, he pursued postdoctoral studies with Professor Vladimir Prelog at ETH Zurich in Switzerland. [13] He worked for nearly a year on alkaloid chemistry in an unpaid position. [10] [17]

University of Liverpool British university

The University of Liverpool is a public university based in the city of Liverpool, England. Founded as a college in 1881, it gained its royal charter in 1903 with the ability to award degrees and is also known to be one of the six original 'red brick' civic universities. It comprises three faculties organised into 35 departments and schools. It is a founding member of the Russell Group, the N8 Group for research collaboration and the university management school is AACSB accredited.

Vladimir Prelog Croatian-Swiss chemist

Vladimir Prelog ForMemRS was a Croatian-Swiss organic chemist who received the 1975 Nobel Prize in chemistry for his research into the stereochemistry of organic molecules and reactions. Prelog was born and grew up in Sarajevo. He lived and worked in Prague, Zagreb and Zürich during his lifetime.

ETH Zurich Swiss Federal Institute of Technology in Zurich

ETH Zurich is a science, technology, engineering and mathematics university in the city of Zürich, Switzerland. Like its sister institution EPFL, it is an integral part of the Swiss Federal Institutes of Technology Domain that is directly subordinate to Switzerland's Federal Department of Economic Affairs, Education and Research. The school was founded by the Swiss Federal Government in 1854 with the stated mission to educate engineers and scientists, serve as a national center of excellence in science and technology and provide a hub for interaction between the scientific community and industry.

During a brief period in 1949, he was unable to find a job in his original home area in the Punjab. [10] He returned to England on a fellowship to work with George Wallace Kenner and Alexander R. Todd on peptides and nucleotides. [17] He stayed in Cambridge from 1950 until 1952.

George Wallace Kenner FRS was a British organic chemist. He was born in Sheffield in 1922, the son of Prof. James Kenner. During his childhood, he went to Didsbury Preparatory School in 1928 and moved to Manchester Grammar School in 1934. He was appointed to the first Heath Harrison Chair of Organic Chemistry at the University of Liverpool 1957–1976. He did his MSc and PhD degrees under Lord Todd at Manchester and Cambridge Universities in UK. He married Jillian Bird in 1951 and they had two daughters both born in Cambridge. He was faculty member at the Cambridge University for 15 years before moving to the University of Liverpool in 1957 as Heath Harrison Professor of Organic Chemistry.

Alexander R. Todd British biochemist

Alexander Robertus Todd, Baron Todd of Trumpington was a Scottish biochemist whose research on the structure and synthesis of nucleotides, nucleosides, and nucleotide coenzymes gained him the Nobel Prize for Chemistry.

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. [13] [18] Khorana was excited by the prospect of starting his own lab, a colleague later recalled. [10] His mentor later said that the Council had few facilities at the time but gave the researcher "all the freedom in the world". [19] His work in British Columbia was on "nucleic acids and synthesis of many important biomolecules" according to the American Chemical Society. [15]

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. [15] [20] He became a professor of biochemistry in 1962 and was named Conrad A. Elvehjem Professor of Life Sciences at Wisconsin–Madison. [21] 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. [15] 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." [22]

He became a US citizen in 1966. [23] Beginning in 1970, Khorana was the Alfred P. Sloan Professor of Biology and Chemistry at the Massachusetts Institute of Technology [24] [13] [25] and later, a member of the Board of Scientific Governors at The Scripps Research Institute. He retired from MIT in 2007. [23]

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.


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. [26]

Their Nobel lecture was delivered on 12 December 1968. [26] Khorana was the first scientist to chemically synthesize oligonucleotides. [27] This achievement, in the 1970s, was also the world's first synthetic gene; in later years, the process has become widespread. [24] Subsequent scientists referred to his research while advancing genome editing with the CRISPR/Cas9 system. [23]

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, [27] as well as methods that anticipated the invention of polymerase chain reaction (PCR). [28] 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. [29] Later, his lab went on to study the structurally related visual pigment known as rhodopsin. [30]

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." [15] [4]

Awards and honors

In addition to sharing the Nobel prize (while he was working at the University of Wisconsin–Madison in the U.S.), [14] Khorana was elected as Foreign Member of the Royal Society (ForMemRS) in 1978. [31] 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, 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: [32] 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. [33] 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. [20]

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. [13]

On 9 January 2018, a Google Doodle celebrated the achievements [34] of Har Gobind Khorana on what would have been his 96th birthday. [35] [36]


Har Gobind Khorana died on 9 November 2011, in Concord, Massachusetts, at the age of 89. [37] His wife, Esther, and daughter, Emily Anne, had died earlier, [15] but Khorana was survived by his other two children. [11] 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." [13]

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". [37]

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Dieter Gerhard Söll is a Sterling Professor of Molecular Biophysics and Biochemistry and Chemistry at the Yale University. He earned his B.S. and Ph.D. from Stuttgart University in 1962 and did his postdoctoral work at University of Wisconsin–Madison from 1962-1965 with Har Gobind Khorana. He was briefly an assistant professor at University of Wisconsin before joining the Yale faculty in 1967 and has been there since. He was named a Sterling Professor in 2006. As a postdoc with Jack Strominger, he identified tRNAs that were involved in peptidoglycan formation leading to the discovery of novel aminoacyl-tRNA functions. He later sequenced the selenocysteine tRNA. His research is centered on the formation of aminoacyl-tRNA and tRNA synthetases. He is a member of the National Academy of Science, fellow of the American Academy of Arts and Sciences, and was named a Guggenheim Fellow in 1972 and 1989 and a Humboldt Fellow in 2000. In addition to his academic work, he has been recognized as a leader in creating research opportunities for minority students notably by spearheading a program to bring students from Tougaloo College to Yale University for summer research in the early 1970s.

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