American Philosophical Society
Member History

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201. Molecular Biology and Biochemistry[X]
61Name:  Dr. Arthur B. Pardee
 Institution:  Dana-Farber Cancer Institute, Harvard University
 Year Elected:  2001
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1921
 Death Date:  February 24, 2019
   
 
Arthur Beck Pardee received his Ph.D. from the California Institute of Technology in 1947. He was an assistant and associate professor at the University of California, Berkeley from 1949-61 and professor of biochemical sciences and Donner Professor of Science at Princeton University from 1961-75. He was a senior postdoctoral fellow at the Pasteur Institute, France from 1957-58 and an American Cancer Society Scholar at the Imperial Cancer Research Fund Laboratory, London, from 1972-73. In 1975 he moved to Cambridge to serve as Professor of Biological Chemistry and Molecular Pharmacology at Harvard Medical School and Chief of the Division of Cell Growth and Regulation at the Dana Farber Cancer Institute. From 1997 on he was professor emeritus at Harvard. Arthur Pardee's early work was in bacterial biochemistry. His studies on growth regulation led to discoveries of repression of gene transcription, feedback inhibition, and allosteric regulation. He next turned to cancer and identified the restriction point, a major regulatory checkpoint that must be bypassed before cells can initiate DNA synthesis. He demonstrated that an unstable protein must be synthesized for a cell to enter S phase, a process defective in cancer cells. He identified cyclin E as the potential restriction point protein, and factor in growth control at the G1/S boundary. An important technical contribution was the development of "differential display," a method that identifies differences in gene expression in various cells and tissues. Dr. Pardee was a recipient of the Paul Lewis Award of the American Chemical Society, the Sir H. A. Krebs Medal, the Rosensteil Medal, the FASEB 3B Award, the CIT Award, the Boehringer-Mannheim Bioanalytica Award, and the Outstanding Alumnus Award of the California Institute of Technology. He was a member of the National Academy of Sciences, the American Academy of Arts & Sciences, the Japanese Biochemical Society, the American Society of Biological Chemists (president, 1980), and the American Association for Cancer Research (president, 1985). He was member of the Cancer Institute Scientific Committee and served on the scientific board of the Worcester Foundation. He was elected a member of the American Philosophical Society in 2001. Arthur Pardee died February 24, 2019 at the age of 97.
 
62Name:  Dr. Alexander Rich
 Institution:  Massachusetts Institute of Technology
 Year Elected:  1980
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1924
 Death Date:  April 27, 2015
   
 
Alexander Rich was a preeminent researcher in structural molecular biology. He joined the faculty of the Massachusetts Institute of Technology in1958, and he had been the William Thompson Sedgwick Professor of Biophysics. Best known for the discovery of left-handed DNA, or Z-DNA, and the three-dimensional structure of transfer RNA, he was also one of the leading workers in the determination of structure and function of Z-DNA binding proteins by x-ray crystallographic and other methods. Dr. Rich received his M.D. from Harvard University Medical School in 1949 and was a founder and director of the pharmaceutical development company Alkermes, Inc. He was awarded the 1995 U.S. National Medal of Science, the 2000 Bower Award for Excellence in Science and the 2009 Welch Award in Chemistry, among other honors, and he has been elected to the National Academy of Sciences and the Institute of Medicine. He was elected a member of the American Philosophical Society in 1980. Dr. Rich died April 27, 2015, at the age of 90, in Boston, Massachusetts.
 
63Name:  Dr. Frederic M. Richards
 Institution:  Yale University
 Year Elected:  1992
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1925
 Death Date:  January 11, 2009
   
 
A pioneer in crystallography and structural biology, Frederic M. Richards has been Sterling Professor Emeritus of Molecular Biophysics and Biochemistry at Yale University since 1991. He received his Ph.D. from Harvard University in 1952 and, aside from a year in Copenhagen with Linderstrom Lang and a year in Cambridge with A.C. Chibnell, he has spent his entire academic career at Yale, chairing the department of molecular biology, biophysics and chemistry from 1969-73. A member of the National Academy of Sciences and the American Academy of Arts & Sciences, Dr. Richards has received honors such as the Pfizer-Paul Lewis Award in Enzyme Chemistry (1965), the Kai Linderstrom-Lang Prize in Protein Chemistry (1978) and the American Society for Biochemistry and Molecular Biology-Merck Award (1988). An intellectual leader, Dr. Richards is admired not only for his meticulous science, which has relevance to many fields, but for his generous, open and warm scientific style.
 
64Name:  Dr. Paul Schimmel
 Institution:  The Scripps Research Institute
 Year Elected:  1999
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1940
   
 
Paul Schimmel is the Ernest and Jean Hahn Professor of Molecular Biology and Chemistry at The Scripps Research Institute. He received his Ph.D. at Massachusetts Institute of Technology and remained on the faculty of MIT until 1997. Dr. Schimmel is the recipient of the Pfizer Award in Enzyme Chemistry from the American Chemical Society and is an elected member of the National Academy of Sciences and American Academy of Arts & Sciences. Dr. Schimmel has contributed extensively to the understanding of the protein-RNA interactions that are the basis of the universal genetic code. He showed how features in small RNA structures are interpreted as specific amino acids referred to by some as a "second genetic code." Most recently he established how RNA structure plays an essential role in enhancing the accuracy of the genetic code by an error correction mechanism. Also, in other work, Nature magazine cited his development of "expressed sequence tags" as one of the four key developments that launched the human genome project. Author or co-author of 400 scientific papers and of a widely used three-volume textbook on biophysical chemistry, Dr. Schimmel has also applied basic biomedical research to human health. For example, his laboratory discovered human proteins active in blood vessel formation, developed them, and brought them to clinical medicine. He holds several patents and is cofounder or founding director of ten biotechnology companies, of which 5 are publicly traded. These companies are developing new therapies for human disease. He was elected to the American Philosophical Society in 1999.
 
65Name:  Dr. Maxine F. Singer
 Institution:  Carnegie Institution of Washington & National Institutes of Health
 Year Elected:  1990
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1931
   
 
Maxine Frank Singer has made major contributions to the biochemistry of nucleic acids and more recently to our knowledge of the mammalian genome structure and organization. She has also served with distinction as chair of the editorial board of the Proceedings of the National Academy of Sciences and as president of the Carnegie Institution, and she is widely recognized as an articulate author and spokesperson for science. After receiving her Ph.D. in biochemistry from Yale University in 1957, Dr. Singer joined the research staff of the National Institutes of Health. She would later serve as chief of the Laboratory of Biochemistry at the National Cancer Institute from 1980-87, where she led fifteen research groups engaged in various biochemical investigations. Dr. Singer's research contributions have ranged over several areas of biochemistry and molecular biology, including chromatin structure, the structure and evolution of defective viruses, and enzymes that work on DNA and its complementary molecule, RNA. In recent years, her foremost contributions have been in studies of a large family of repeated DNA sequences called LINES - sequences interspersed many times in mammalian DNA. She and her co-workers have been especially interested in the LINE-1 sequence, which is repeated thousands of times in human DNA. LINE-1, she early concluded, is capable of insertion into new places on chromosomal DNA, and researchers elsewhere later found that LINE-1 insertions into a gene whose product is required for blood clotting are associated with cases of hemophilia. Believing that the mechanism of LINE-1 transposition might have broad significance for understanding genetic diseases, Dr. Singer and her colleagues have concentrated their experiments on learning how LINE-1 elements move. Throughout her career, Dr. Singer has assumed leading roles in influencing and refining the nation's science policy. In 1988 she became President of the Carnegie Institution of Washington, where she led the biologists, astronomers and earth scientists who make up the Institution's six scientific departments. Dr. Singer is presently President Emerita of the Carnegie Institution while also retaining her association with the National Cancer Institute as Scientist Emerita. Her several awards for public service include the Distinguished Presidential Rank Award (1988) and the National Medal of Science (1992), the nation's highest scientific honor bestowed by the President of the United States, "for her outstanding scientific accomplishments and her deep concern for the societal responsibility of the scientist." Most recently, she was honored with the 2007 Public Welfare Medal, the National Academy of Sciences' most prestigious award recognizing extraordinary use of science for the public good.
 
66Name:  Dr. Emil L. Smith
 Institution:  University of California, Los Angeles
 Year Elected:  1973
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1911
 Death Date:  May 31, 2009
   
 
Emil L. Smith is Professor of Biological Chemistry Emeritus at the University of California, Los Angeles, where he has served on the faculty of the School of Medicine since 1963. One of the world's leading authorities on the amino acid sequences of proteins and on biochemical evolution, his research has dealt with photosynthesis; chlorophyll; physiology of the visual process; proteolytic enzymes; glycoproteins; cytochrome; histones; and glutamate dehydrogenases. An outstanding leader in biochemical research and a man of broad scientific interests, Dr. Smith earned his Ph.D. from Columbia University in 1937. He subsequently worked as a Guggenheim Fellow at Cambridge and Yale and as a research associate at the Rockefeller Institute (1940-42) and as a biophysicist at E.R. Squibb & Company's Biological Laboratories (1942-46) before joining the faculty of the University of Utah (1946-63). From 1959-62 Dr. Smith also served as chairman of the United States National Committee on Biochemistry. A member of the National Academy of Sciences, he is the author (with A. White, P. Handler and others) of Principles of Biochemistry, for seven editions (1954-83) one of the leading textbooks in the field.
 
67Name:  Dr. Alexander S. Spirin
 Institution:  Moscow State University & Russian Academy of Sciences
 Year Elected:  1997
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  International
 Living? :   Deceased
 Birth Date:  1931
 Death Date:  December 30, 2020
   
 
Alexander S. Spirin was a world-class Russian scientist who has contributed much that is ingenious and original to our understanding of the structure and function of ribosomes - the intricate molecular machines that synthesize the proteins of cells. He has provided fascinating insight into the interplay between various types of ribonucleic acids and proteins that make up these machines. He has done this by taking ribosomes apart and then sucessfully reassembling them: a major achievement. Currently Professor of Biochemistry and Molecular Biology at Moscow State University and the Director of the Institute of Protein Research at the Russian Academy of Sciences, Dr. Spirin has created a body of work that is important, elegant and internationally recognized. He died on December 30, 2021.
 
68Name:  Dr. Donald F. Steiner
 Institution:  University of Chicago; Howard Hughes Medical Institute
 Year Elected:  2004
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1930
 Death Date:  November 11, 2014
   
 
Donald F. Steiner was born in Lima, Ohio in 1930. He received his M.D. degree from the University of Chicago in 1956 and had a distinguished career at the university as professor of biochemistry (1968-70); A. N. Pritzker Professor of Biochemistry and Medicine (1970-84); chairman of the department of biochemistry (1973-79); director of the Diabetes-Endocrinology Center (1974-78); associate director of the Diabetes and Research Training Center (1977-81); and A. N. Pritzker Distinguished Service Professor of Biochemistry & Molecular Biology and Medicine (1984-2014). He has also been a senior investigator in the Howard Hughes Medical Institute (1985-2006) and director or co-director of the University of Chicago Diabetes and Research Training Center. In 1967 Dr. Steiner discovered proinsulin, the single-chain precursor of insulin. He purified it and studied its structure, properties, biosynthesis, and cell biology, demonstrating its intracellular conversion into insulin and the cosecreted C-peptide. With Dr. A. H. Rubenstein, radioimmunoassays were developed for proinsulin and C-peptide in serum, which have been widely applied in diabetes research. Dr. Steiner's pioneering studies thus opened the now very broad field of precursor protein processing, leading to the identification of many other proproteins and more recently to the discovery of the mammalian proprotein convertase family of cellular processing endoproteases. His laboratory also first demonstrated receptor-mediated uptake and degradation of insulin. His discoveries have strongly influenced insulin and islet cell research, ranging from the commercial production of human insulin for diabetes therapy to the evolution of insulin and insulin-like growth factors (IGFs). The recipient of honors including the Gairdner Award (1971), Wolf Foundation Prize in Medicine (1985) and the Endocrine Society's Fred C. Koch Award (1990), Dr. Steiner was a member of the National Academy of Sciences and the American Academy of Arts & Sciences. He was elected a member ofthe American Philosophical Society in 2004. Donald Steiner died November 11, 2014, at age 84 at his home in Chicago, Illinois.
 
69Name:  Dr. Joan A. Steitz
 Institution:  Yale University & Howard Hughes Medical Institute
 Year Elected:  1992
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1941
   
 
Joan A. Steitz is Sterling Professor of Molecular Biophysics and Biochemistry at Yale University as well as an investigator at the Howard Hughes Medical Institute. She has made important research contributions to the study of the role of small nuclear RNAs (snRNAs) in eukaryotic cells, and she is credited with discovering that complexes containing these snRNAs reacted specifically with sera from autoimmune patients. She then recognized that sequences in one of these snRNAs were complementary to sequences important in splicing of RNA in the nucleus of cells. This hypothesis proved correct and played a critical role in directing the field. Dr. Steitz is the recipient of prestigious awards such as the Eli Lilly Award in Biological Chemistry (1982), the National Medal of Science (1987), the Dickson Prize for Science (1989), the Cristopher Columbus Discovery Award in Biomedical Research (1992), the Lewis S. Rosenstiel Award (2002), the Albany Medical Center Prize in Medicine and Biomedical Research (2008), the Connecticut Medal of Science (2015), the William Clyde DeVane Medal (2016), and the Wolf Prize in Medicine (2021). A graduate of Antioch College (B.S., 1963) and Harvard University (Ph.D., 1967), she was elected to the membership of the American Academy of Arts & Sciences in 1982 and the National Academy of Sciences in 1983.
 
70Name:  Dr. Jack W. Szostak
 Institution:  Howard Hughes Medical Institute; Harvard Medical School; Massachusetts General Hospital
 Year Elected:  2012
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1952
   
 
Dr. Jack W. Szostak is an Investigator of the Howard Hughes Medical Institute, Professor of Genetics at Harvard Medical School, and the Alex Rich Distinguished Investigator in the Dept. of Molecular Biology and the Center for Computational and Integrative Biology at Massachusetts General Hospital. Dr. Szostak is a member of the National Academy of Sciences, and a Fellow of the New York Academy of Sciences, the American Academy of Arts and Sciences, and the American Association for the Advancement of Science. Dr. Szostak’s early research was on the genetics and biochemistry of DNA recombination, which led to the double-strand-break repair model for meiotic recombination. At the same time Dr. Szostak made fundamental contributions to our understanding of telomere structure and function, and the role of telomere maintenance in preventing cellular senescence. For this work Dr. Szostak shared, with Drs. Elizabeth Blackburn and Carol Greider, the 2006 Albert Lasker Basic Medical Research Award and the 2009 Nobel Prize in Physiology or Medicine. In the 1990s Dr. Szostak and his colleagues developed in vitro selection as a tool for the isolation of rare functional RNA, DNA and protein molecules from large pools of random sequences. His laboratory has used in vitro selection and directed evolution to isolate and characterize numerous nucleic acid sequences with specific ligand binding and catalytic properties. For this work, Dr. Szostak was awarded, along with Dr. Gerald Joyce, the 1994 National Academy of Sciences Award in Molecular Biology and the 1997 Sigrist Prize from the University of Bern. In 2000, Dr. Szostak was awarded the Medal of the Genetics Society of America, and in 2008 Dr. Szostak received the H.P. Heineken Prize in Biophysics and Biochemistry. Dr. Szostak’s current research interests are in the laboratory synthesis of self-replicating systems and the origin of life.
 
71Name:  Dr. Paul Talalay
 Institution:  Johns Hopkins University
 Year Elected:  1990
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1923
 Death Date:  March 10, 2019
   
 
Paul Talalay, M.D. was John Jacob Abel Distinguished Service Professor of Pharmacology and Molecular Sciences at Johns Hopkins University School of Medicine. He held an S.B. degree in biophysics from M.I.T. and an M.D. degree from Yale. Following surgical training at Massachusetts General Hospital, he moved to the University of Chicago, rising to the academic ranks of Professor of Biochemistry, Professor of Medicine, and Professor in the Ben May Laboratory for Cancer Research. After serving for 12 years as Director of the Department of Pharmacology at Johns Hopkins Medical School, he relinquished this position to devote himself full time to research. Dr. Talalay devoted his career to cancer research. For more than 2 decades he was involved in devising strategies for chemoprotection against the risk of cancer, a field in which he is recognized as a pioneer. His efforts focused on achieving protection by raising the enzymes concerned with the detoxication of carcinogens. Analysis of the chemistry and the molecular biology of boosting enzymes of detoxication led him and his colleagues to devise simple cell culture methods for detecting chemical and especially dietary (Phyto)chemicals that raise these enzymes. This work led to the isolation of sulforaphane as the most potent inducer of protective enzymes in broccoli. These findings, published in the Proceedings of the National Academy of Sciences, attracted world-wide attention, and led to the organization of the Brassica Chemoprotection Laboratory at Johns Hopkins. This unique laboratory is exclusively dedicated to identifying edible plants that are particularly rich in protective enzyme-inducer activity. Dr. Talalay's honors, in addition to his appointment as a University Distinguished Service Professor, included appointment to one of the first lifetime professorships of the American Cancer Society and membership in the National Academy of Sciences and the American Academy of Arts & Sciences. He published more than two hundred papers in internationally respected scientific journals. He received an honorary D.Sc. degree from Acadia University. The M.D.-Ph.D. Student Library at Johns Hopkins has been named in his honor. He was elected a member of the American Philosophical Society in 1990. Paul Talalay died March 10, 2019 in Baltimore, MD at the age of 95.
 
72Name:  Dr. Rudolf K. Thauer
 Institution:  Max Planck Institute for Terrestrial Microbiology; Philipps University Marburg
 Year Elected:  2018
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  International
 Living? :   Living
 Birth Date:  1939
   
 
Born 1939 in Frankfurt, Germany, I went to school in Wetter, Landshut, Philadelphia, USA (1947-1951) and Bad Nauheim and then studied Medicine and Biochemistry at the Universities of Frankfurt, Tubingen and Freiburg, where I ended my studies 1968 with a PhD in Biochemistry and a Thesis on the "Energy Metabolism of Clostridium kluyveri." Ever since then my scientific interest remained focused on how strictly anaerobic microorganisms conserve energy. Discoveries made were amongst others that carbon monoxide is an intermediate in autotrophic CO2 fixation via the Wood-Ljungdahl pathway and that the trace element nickel is required by many anaerobes as cofactor of carbon monoxide dehydrogenase, hydrogenases and methyl-coenzyme M reductase. The latter enzyme has a nickel tetrapyrrole as prosthetic group and catalyzes both methane formation and methane oxidation in Archaea. After a short postdoc in 1971 with Harland Wood at Case Western University Chicago, I was appointed in 1972 Associate Professor for Biochemistry at the Ruhr University in Bochum, in 1976 Full Professor for Microbiology at the Philipps University Marburg and in 1991 Founding Director of the Max Planck Institute for Terrestrial Microbiology in Marburg. After my retirement as Director at the end of 2007 I continued research at the Max Planck Institute that led together with Wolfgang Buckel to the discovery of flavin-based electron bifurcation that changed our understanding how most anaerobes conserve energy.
 
73Name:  Dr. Robert Tjian
 Institution:  University of California, Berkeley; Howard Hughes Medical Institute
 Year Elected:  2009
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Living
 Birth Date:  1949
   
 
Robert Tjian served as president of the Howard Hughes Medical Institute in April 2009 through the end of 2016 and then returned to his lab at the University of California, Berkeley. Trained as a biochemist, he has made major contributions to the understanding of how genes work during three decades on the faculty of the University of California, Berkeley. He was named an HHMI investigator in 1987. Tjian studies the biochemical steps involved in controlling how genes are turned on and off, key steps in the process of decoding the human genome. He discovered proteins called transcription factors that bind to specific sections of DNA and play a critical role in controlling how genetic information is transcribed and translated into the thousands of biomolecules that keep cells, tissues, and organisms alive. Tjian's laboratory has illuminated the relationship between disruptions in the process of transcription and human diseases such as cancer, diabetes, and Huntington's. More recently, he has begun studying how transcription factors control the differentiation of embryonic stem cells into muscle, liver, and neurons. Tjian, 59, was born in Hong Kong, the youngest of nine children. His family fled China before the Communist Revolution and eventually settled in New Jersey. Known as a voracious consumer of scientific information and data, Tjian famously talked his way into the biochemistry laboratory of the late Daniel Koshland as a Berkeley undergraduate—even though he had never taken a single course in the subject. Tjian went on to receive a bachelor's degree in biochemistry from Berkeley in 1971 and a Ph.D. from Harvard University in 1976. After completing a postdoctoral fellowship at the Cold Spring Harbor Laboratory with James Watson, he joined the Berkeley faculty in 1979. At Berkeley, Tjian assumed a variety of leadership roles, including spearheading a major campus initiative to support and implement new paradigms for bioscience teaching and research. He served as the Director of the Berkeley Stem Cell Center, and the Faculty Director of the Li Ka Shing Center for Biomedical and Health Sciences. He is a member of the National Academy of Sciences and has received many awards honoring his scientific contributions, including the Alfred P. Sloan Prize from the General Motors Cancer Research Foundation and the Louisa Gross Horwitz Prize from Columbia University. He was named California Scientist of the Year in 1994. As president of the Institute, Tjian remains an active scientist. His small laboratory group at HHMI's Janelia Farm Research Campus is focused on the development of new approaches to image biochemical activities in single living cells. He will also maintain a research laboratory at UC Berkeley, where he is a professor of biochemistry and molecular biology. Tjian and his wife, Claudia, an attorney, have two daughters.
 
74Name:  Dr. Don Craig Wiley
 Institution:  Harvard & Howard Hughes Medical Institute
 Year Elected:  1996
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1944
 Death Date:  November 16, 2001
   
75Name:  Dr. Bernhard Witkop
 Institution:  National Institutes of Health
 Year Elected:  1999
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1917
 Death Date:  November 22, 2010
   
 
Bernhard Witkop was a distinguished organic chemist who had made major contributions to biochemistry, pharmacology, and medical sciences. His research involved major contributions to the chemistry of familiar-sounding substances or processes, such as the isolation from the Colombian tree frog of the toxin batrachotoxin, a godsend to electrophysiologists. He made important contributions to research on antiviral agents, interferon, dopamine, genetically engineered proteins and metabolic pathways. His cyanogen bromide cleavage reaction made possible the production of the first 100 Kg of engineered insulin by Eli Lilly. In Dr. Witkop's paper "Mind Over Matter" he assumes "the uneasy role of the scientist as philosopher" and presents a scholarly and profound contribution on this topic so central to most philosophers. Later in his life Dr. Witkop dedicated efforts to historical biography. He worked at the National Institutes of Health since 1987, where he is Scholar Emeritus. A native of Germany, Dr. Witkop held Ph.D. (1940) and Sc.D. (1946) degrees from the University of Munich.
 
76Name:  Dr. Paul C. Zamecnik
 Institution:  Massachusetts General Hospital, Harvard Medical School
 Year Elected:  2006
 Class:  2. Biological Sciences
 Subdivision:  201. Molecular Biology and Biochemistry
 Residency:  Resident
 Living? :   Deceased
 Birth Date:  1912
 Death Date:  October 27, 2009
   
 
Paul Zamecnik is a senior scientist at Massachusetts General Hospital and Professor Emeritus at Harvard Medical School. He has been affiliated with both institutions for over fifty years and received his M.D. from Harvard Medical School in 1936. Dr. Zamecnik's first great scientific contribution was elucidating important aspects of the biochemistry of protein synthesis. He showed that ATP is necessary for peptide bond formation, which therefore is not a reversal of proteolysis; in the cell free system, he devised the ribosome is the site of these reactions; and tRNAs translate sequences of DNA to protein. Early, he performed RNA sequencing that showed 3'-poly A in Rous sarcoma virus RNA, and a prior sequence identical to that at the 5' end, indicating circular structure. His second greatest contribution was the conception of competitive antisense nucleotides. These blocked virus replication by inhibition of translation. He demonstrated the antisense effect with hemoglobin protein synthesizing cells showing that this depends on the ability of deoxynucleotides to enter intact cells and on Watson-Crick base pairing. He has also applied the concept to medicine, targeting the tuberculosis bacterium and the defective cystic fibrosis gene. A three-time winner of the John Collins Warren Triennial Prize, (1946, 1950, 1999) as well as the Presidential Medal of Science (1991), the Lasker Award (1995) and the Institute of Virology's Lifetime Achievement Award (2004), Dr. Zamecnik was elected to the membership of the American Academy of Arts & Sciences in 1954, the National Academy of Sciences in 1968 and the American Philosophical Society in 2006.
 
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