Holley, Robert William

born Jan. 28, 1922, Urbana, Ill., U.S.

died Feb. 11, 1993, Los Gatos, Calif.

U.S. biochemist.

He received his Ph.D. from Cornell University. Holley and others showed that transfer RNA was involved in the assembly of amino acids into proteins. He was the first to determine the sequence of nucleotides in a nucleic acid, a process that required digesting the molecule with enzymes, identifying the pieces, and then figuring out how they fit together. It has since been shown that all transfer RNA has a similar structure. He shared a 1968 Nobel Prize with Marshall Warren Nirenberg and Har Gobind Khorana.

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▪ 1994

      U.S. biochemist (b. Jan. 28, 1922, Urbana, Ill.—d. Feb. 11, 1993, Los Gatos, Calif.), shared the 1968 Nobel Prize for Physiology or Medicine with H. Gobind Khorana and Marshall W. Nirenberg; the three scientists independently conducted research that helped to decipher the genetic code chemically and explain how the genetic information stored in the DNA of a cell controls the synthesis of proteins, the building blocks of cells. Holley, who began his painstaking work in 1956 while at the U.S. Department of Agriculture's Plant, Soil and Nutrition Laboratory at Cornell University, Ithaca, N.Y., determined the structure of alanine transfer RNA by purifying small amounts of the RNA isolated from more than 135 kg (300 lb) of baker's yeast. In 1965 Holley reported that "the complete nucleotide sequence of an alanine transfer RNA, isolated from yeast, has been determined. This is the first nucleic acid for which the structure is known." After earning a Ph.D. in organic chemistry (1947) from Cornell, Holley became associated with the university's state and federal agricultural stations and taught biochemistry and molecular biology (1962-66), serving at chairman of the biochemistry department from 1965 to 1966. In 1968 he became a resident fellow at the Salk Institute for Biological Studies in La Jolla, Calif., where he remained until his death. There he studied both the normal and the abnormal functions of the growth of cells in mammals, primarily focusing on the timing of cell division. The latter was crucial to understanding the growth of cancer and aided in diagnosis and treatment of that disease and others. Holley was also the recipient of the prestigious Lasker Award (1965) and of an award in molecular biology (1967) from the National Academy of Sciences.

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▪ American biochemist

born Jan. 28, 1922, Urbana, Ill., U.S.

died Feb. 11, 1993, Los Gatos, Calif.

      American biochemist who shared the Nobel Prize in Physiology or Medicine in 1968 with Marshall Warren Nirenberg (Nirenberg, Marshall Warren) and Har Gobind Khorana (Khorana, Har Gobind). Their research helped explain how the genetic code controls the synthesis of proteins.

      Holley obtained his Ph.D. in organic chemistry from Cornell University, Ithaca, N.Y., in 1947. He investigated a variety of biochemical questions at the state and federal agricultural experiment stations at Cornell (1948–64). He began his research on RNA after spending a year studying with James F. Bonner at the California Institute of Technology (1955–56).

      By 1960 Holley and others had shown that small molecules of ribonucleic acids, called transfer RNAs, were involved in the assembly of amino acids into proteins. Holley and his collaborators developed techniques to separate the different transfer RNAs from the mixture in the cell. By 1965 he had determined the composition of the transfer RNA that incorporates the amino acid alanine into protein molecules. This feat—the first determination of the sequence of nucleotides in a nucleic acid—required digesting the molecule with enzymes, identifying the pieces, then figuring out how they fit together. It has since been shown that all transfer RNAs have similar structures.

      In 1968 Holley became a resident fellow at the Salk Institute for Biological Studies in La Jolla, Calif. He also became an adjunct professor at the University of California, San Diego, in the following year.

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