The following sections of this BookRags Literature Study Guide is offprint from Gale's For Students Series: Presenting Analysis, Context, and Criticism on Commonly Studied Works: Introduction, Author Biography, Plot Summary, Characters, Themes, Style, Historical Context, Critical Overview, Criticism and Critical Essays, Media Adaptations, Topics for Further Study, Compare & Contrast, What Do I Read Next?, For Further Study, and Sources.
(c)1998-2002; (c)2002 by Gale. Gale is an imprint of The Gale Group, Inc., a division of Thomson Learning, Inc. Gale and Design and Thomson Learning are trademarks used herein under license.
The following sections, if they exist, are offprint from Beacham's Encyclopedia of Popular Fiction: "Social Concerns", "Thematic Overview", "Techniques", "Literary Precedents", "Key Questions", "Related Titles", "Adaptations", "Related Web Sites". (c)1994-2005, by Walton Beacham.
The following sections, if they exist, are offprint from Beacham's Guide to Literature for Young Adults: "About the Author", "Overview", "Setting", "Literary Qualities", "Social Sensitivity", "Topics for Discussion", "Ideas for Reports and Papers". (c)1994-2005, by Walton Beacham.
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Robert William Holley is best known for his 1962-1965 work in chemically isolating pure strands of transfer ribonucleic acid (tRNA), which retrieves specific amino acids for assembly by the cell's ribosomes into proteins. He also determined the complete sequence of one RNA strand. For this work, he shared the 1968 Nobel Prize for physiology or medicine with Har Gobind Khorana and Marshall Nirenberg.
Holley was born in Urbana, Illinois, where his parents were teachers, and received his bachelor's degree from the University of Illinois. He obtained his Ph.D. from Cornell University and spent almost his entire career there, in 1964 becoming professor of biochemistry and molecular biology.
Working with yeast RNA, Holley used techniques developed for proteins by Frederick Sanger, including chemical fragmenting, cutting with restriction enzymes, and separating by ion-exchange chromatography and paper electrophoresis. He used snake venom for the first cuts on long strands. After deciphering the base sequences on the fragments, he reconstructed the sequence of the entire strand.
Following this, Holley studied the relationship between messenger RNA (mRNA), the first copy of a DNA instruction, and tRNA. He also partially identified the three-dimensional structure of RNA and showed that tRNA was a cloverleaf, rather than a helix as deoxyribonucleic acid (DNA) is.