CURRICULUM VITAE
HAROLD ELIOT VARMUS, M.D.
Date of Birth: December 18, 1939 Place of Birth: Oceanside, New York Email Address: [email protected] Spouse: Constance Casey Children: Jacob Carey Varmus, Christopher Isaac Varmus Education Freeport High School, Freeport, New York Amherst College, Amherst, Massachusetts B.A. magna cum laude 1961 Harvard University, Cambridge, Massachusetts M.A. 1962 Columbia University, New York City, New York M.D. 1966 Training Intern in Medicine, Presbyterian Hospital, New York, 1966-1967. Assistant Resident in Medicine, Presbyterian Hospital, New York City, 1967-1968. Clinical Associate, National Institute of Arthritis and Metabolic Diseases, Bethesda, Maryland,
1968-1970. Postdoctoral Fellow, Department of Microbiology, University of California Medical Center,
San Francisco, California, 1970-1972. Appointments Lecturer, Department of Microbiology, University of California Medical Center, San Francisco,
California, 1970-1972. Assistant Professor in Residence, Department of Microbiology, University of California Medical
Center, San Francisco, California, 1972-1974. Associate Professor, Department of Microbiology, University of California Medical Center, San
Francisco, California, 1974-1979. Professor, Department of Microbiology and Immunology, University of California Medical Center,
San Francisco, California, 1979-1993, Department of Biochemistry and Biophysics, 1982-1993; on leave, 1993-March 2000.
American Cancer Society Professor of Molecular Virology, 1984-1993. Chief, Varmus Lab, Division of Basic Sciences, National Cancer Institute, 1993-1999. Director, National Institutes of Health, 1993-1999. President and Chief Executive Officer, Memorial Sloan-Kettering Cancer Center, 2000-2010. Acting Chairman, Sloan-Kettering Institute, 2000-2002.
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Member, Cancer Biology and Genetics Program, SKI, 2000-2010. Attending Physician- Molecular Biology, Department of Medicine, Memorial Hospital, 2000-2010. Acting Director (2000-2002) and Professor - Sloan-Kettering Division, Weill Graduate School of Medical Sciences, Cornell University - Cell Biology & Genetics Program, 2000-2010. Head, Varmus Lab, SKI, 2000-2010. Director, National Cancer Institute, National Institutes of Health, 2010-2015. Lewis Thomas University Professor Of Medicine, Weill Cornell Medical College, 2015-present. Military Service U.S. Public Health Service, Surgeon, 1968-1970. Societies Phi Beta Kappa; Alpha Omega Alpha; American Association for the Advancement of Science; American Society for Cell Biology, American Society for Microbiology; American Society for Biochemistry and Molecular Biology; American Society for Virology; National Academy of Sciences (1984); American Academy of Arts and Sciences (1988); Institute of Medicine (1991); American Association of Physicians (1994); American Philosophical Society (1994); The Royal Society, Foreign Member (2005). Fellowships and Awards Woodrow Wilson Fellow, 1961-1962 International Fellow of Columbia University, 1963-1964 Smith, Kline, and French Foreign Fellow (Bareilly, India), 1966 Senior Dernham Fellow, California Division, American Cancer Society, 1970-1972 USPHS Research Career Development Awardee, 1972-1977 Josiah Macy Scholar, Imperial Cancer Research Fund, London, 1978-1979 California Scientist of the Year, 1982 Albert Lasker Basic Medical Research Award, 1982 Passano Foundation Award, 1983 Armand Hammer Cancer Prize, 1984 Alfred P. Sloan Prize, General Motors Cancer Research Foundation, 1984 Simon M. Shubitz Cancer Prize, 1984 Gairdner Foundation International Award, 1984 D. Sc. (Hon.), Amherst College, 1984 Joseph Mather Smith Award, Columbia College of Physicians & Surgeons, 1986 American College of Physicians Award, 1987 Gold Medal, Alumni Association of the College of Physicians & Surgeons, 1989 Nobel Prize in Physiology or Medicine, 1989 D. Sc. (Hon.), Columbia University, 1990 University of California, San Francisco, Medal, 1990 D. Sc. (Hon), University of Maryland, Baltimore County, 1994 D. Sc. (Hon.), Harvard University, 1996 D. Sc. (Hon.), Dartmouth College, 1997 Deans Medal, Johns Hopkins University, School of Public Health, 1998
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D. Sc. (Hon.), Princeton University, 1999 D. Sc. (Hon.), Wesleyan University, 2000 D. Sc. (Hon.), Yale University, 2001 D. Sc. (Hon.), Bard College, 2001 D. Sc. (Hon.), Mt. Sinai School of Medicine, 2001 Vannevar Bush Award, National Science Foundation, 2001 Barnard Medal of Distinction, Barnard College, Columbia University, 2002 National Medal of Science, 2002 D. Sc. (Hon.), University of Rochester, 2004 D. Sc. (Hon.), University of Miami, 2004 WIRED Magazine’s Rave Award, 2004 Jill Rose Award, The Breast Cancer Research Foundation, 2004 D. Sc. (Hon.), SUNY Downstate’s College of Medicine and Graduate Studies, 2005 America’s Best Leaders Award, U.S. News and World Report and the Center for Public Leadership, 2007 Henry G. Friesen International Prize in Health Research, Friends of the Canadian Institutes of Health Research and Canadian Academy of Health Sciences, 2008 Thomas C. Schelling Award, John F. Kennedy School of Government, Harvard University, 2009 Phi Beta Kappa Book Award in Science, Phi Beta Kappa Society, 2009 D. Sc. (Hon.), Rensselaer Polytechnic Institute, 2010 Honorary Degree, University of Massachusetts Medical School, 2010 D.Sc. (Hon.), The Rockefeller University, 2010 Double Helix Medal, Cold Spring Harbor Laboratory, 2011 D. Sc. (Hon.), Louis V. Gerstner, Jr. Graduate School of Biomedical Sciences, MSKCC, 2012 Memorial Sloan-Kettering Medal for Outstanding Contributions to Biomedical Research, 2012 Pupin Medal, Columbia University & The Columbia Engineering Alumni Association, 2012 Glenn T. Seaborg Medal, UCLA, 2012 Doctor of Letters in Medicine (Hon.), Baylor College of Medicine, 2013 D. Sc. (Hon.), University of Cambridge, 2013 The John P. McGovern Award, Cosmos Club Foundation, 2013 International Agency for Research on Cancer (IARC) Medal of Honor, 2014 Doctor of Humane Letters (Hon.), Macaulay Honors College, The City University of NY, 2014 Professional Activities Scientific Journals Associate Editor, Cell, 1974-1978, 1979-1993 Associate Editor, Virology, 1974-1984 Editor, Molecular and Cellular Biology, 1984-1988 Associate Editor, Genes and Development, 1986-1993 Editorial Board, Trends in Genetics, 1989-1993 Editorial Board, Proceedings of National Academy of Sciences, 1987-1993 Editorial Board, Cancer Surveys, 1985-1993 Editorial Board, Current Biology, 1992-1993 Associate Editor, Molecular Biology of the Cell, 2000-2004 Board of Trustees, BioMed Central, 2000-2008 Editorial Board, Journal of Biology, 2002-2007
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Senior Editor, The Journal of Clinical Investigation, 2002 – 2007 Non-Profit Advisory and Review Boards Special Grants Committee, California Division of the American Cancer Society, 1973-1976 Breast Cancer Virus Working Group, Virus Cancer Program, National Cancer Institute, 1973-1974 Virology Study Section, National Institutes of Health, 1976-1980 Chairman, Retrovirus Study Group, International Committee on Taxonomy of Viruses, 1981-1987 Personnel Committee, American Cancer Society, 1982-1986 Board of Scientific Counselors, Division of Cancer Biology and Diagnosis, National Cancer Institute, 1983-1987 Research Fellowship Committee, California Division, American Cancer Society, 1986-1989 Human Gene Therapy Subcommittee, Recombinant DNA Advisory Committee, NIH, 1986-1989 Chairman, Animal Cells and Viruses Gordon Conference, 1987 Searle Scholars Review Committee, 1988-1992 Genetics Scientific Review Board, Howard Hughes Medical Institute, 1990-1993 Advisor, Congressional Caucus for Biomedical Research, 1990-1993 Member, the Joint Steering Committee for Public Policy of the Biomedical Societies, 1991-1993. Scientific Review Board, Sloan-Kettering Institute, 1991-1993 Visiting Committee, Department of Cell and Developmental Biology, Harvard University,
1991-1992 Chairman, Board on Biology, National Research Council, 1991-1993 Co-Chairman, New Delegation for Biomedical Research, 1991-1993 1994-1999: none Chair, Coalition for the Life Sciences, formerly the Joint Steering Committee for Public Policy of the Biomedical Societies, 2000-2008 Commissioner, World Health Organization’s Commission on Macroeconomic and Health (CMH), 2000-2002 Co-Chair, Committee on Agricultural Biotechnology, Health, and the Environment,
National Research Council, 2000-2002 PubMed Central National Advisory Committee, NCBI, NLM, NIH, 2000-2003 Board of Directors, Blanchette Rockefeller Neurosciences Institute, 2000-2003 Pasteur Institute, Chairman - External Scientific and Strategic Committee (CEOSS), 2000-2004 Scientific Advisory Committee, Imperial Cancer Research Fund, 2000-2002 AIDS Vaccine Research Working Group, NIAID, NIH, 2000-2002 Corporation of The Jackson Laboratory, 2001-2007 Council on Foreign Relations, 2001-Present; CFR Task Force on America’s Reponse to Terrorism and Public Health and Bio-Terrorism Sub-Group, 2001-2002; Independent Task Force on Homeland Security Imperatives, 2002 Coordinating Committee, The Cyprus Institute, 2001-2007 Public Policy Committee, The American Society for Cell Biology, 2001-2003 Distinguished Counsellor to the Board, The New York Botanical Garden, 2001-2010 Lasker Prize Jury, 2001-2010
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Medical Advisory Board, Howard Hughes Medical Institute, 2001-2003 External Review Committee, Alliance for Cell Signaling, NIGMS, NIH, 2001-2002 External Advisory Committee for the Life Sciences, Cornell University, 2001- 2007 Co-Founder, 2001, and Chairman, Board of Directors, Public Library of Science, 2002-2010 Board, Science Institutes Group (Millennium Science Initiative), 2002-2010 Board of Trustees, Columbia University, 2002-2005 Earth Institute Advisory Board, Columbia University, 2002 – 2008 Advisory Board, Millennium Project of the United Nations, 2002 – 2006 President’s Interntional Advisory Committee, Canadian Institutes of Health Research, 2003 – 2006 Chair, Scientific Board, Grand Challenges in Global Health Initiative, Bill & Melinda Gates Foundation and Foundation for the NIH, 2003-2008 Patron, Multilateral Initiative on Malaria, 2003-2010 Mayoral Advisory Task Force on Stimulating Economic Growth and Diversification in New York City Economy, The Partnership for New York City and NYC Economic Development Corporation, 2003-2006 CEO’s Council, New York Academy of Sciences, 2003-2010 Ad-Hoc Subcommittee on a National Advanced Technology Incentive for Cancer, National Cancer Advisory Board, 2003-2006 General Motors Cancer Research Award Committee, 2004-2006 Board of Trustees, Italian Institute of Technology, 2004 – 2005 Member, National Academies Elementary Particle Physics in the 21st Century Committee, 2004 – 2006 Board of Scientific Counselors, The Broad Institute, 2005-2010 Study Group on Science and the Media, American Academy of Arts and Sciences, 2005-2007 Medical Advisory Board, New York Stem Cell Foundation, 2005-2008 Scientific Advisor, World Science Festival, 2005-2010 Advisory Board, DefCon: Campaign to Defend the Constitution, 2005-2007 Council of Trustees, Cancer Research UK, 2006-2008; Council Research Strategy Committee, 2006-2008 Board of Advisors, E.O. Wilson Biodiversity Foundation, 2006-2008 Member, American Meteorological Society Advisory Committee, 2006-2010 Review Committee, Trieste Science Prize, Academy of Sciences for the Developing World, 2006 Scientific Advisory Board, Center for HIV/AIDS Vaccine Immunology (CHAVI), 2006-2008 Member, Transportation Alternatives Advisory Council, 2007-2010 Member, Canary Foundation Strategic Advisory Group, 2007-2010 Co-Chair, Center for an Urban Future’s “Building New York’s Innovation Economy: Leveraging Science & Technology Research for Economic Development” Study Advisory Board, 2007- 2008
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Honorary Board Member, Annette Urso Rickel Foundation, Inc., 2007-2010 Commissioner, Empire State Stem Cell Commission, Funding Committee, 2007- 2008 Chair, Program Advisory Panel of the Bill & Melinda Gates Foundation Global Health Program, 2007-2010 Member, Alzheimer’s Study Group, 2007-2009 Member, Council of the Enterprise, Global HIV Vaccine Enterprise, 2008 Board of Advisors, Scientists & Engineers for America, 2008 Co-Chair, Committee on the U.S. Commitment to Global Health, Institute of Medicine, 2008-2009 Member, Study Group on Presidential Science and Technology Advisory Assets, Center for the Study of the Presidency, 2008 Member, INSERM Evaluation Committee, French Agency for Evaluation of Research and Higher Education, 2008 Member, Founders’ Assembly, Foundation Lindau Nobel Laureate Meetings, 2008-2010 Co-Chair, President’s Council of Advisors on Science and Technology, 2009-2010 Industrial Advisory Groups Scientific Consultant, Chiron Corporation, 1982-1987 Scientific Advisory Board, Merck Corporation, 1985-1988 Scientific Advisory Board, Gilead Corporation, 1988-1993 Chairman, Scientific Advisory Board, Onyx Corporation, 1992-1993 UCSF Service (Partial List) Advisory Council, Medical Scientist Training Program, UCSF, 1977-1993 Steering Committee, Department of Microbiology and Immunology, UCSF, 1982-1993 Vice-Chairman, Board of Overseers, UCSF Biomolecular Resource Laboratory, 1986-1993 Scientific Advisory Board, Gladstone Institute for Virology and Immunology (UCSF AIDS
Research Center), 1993 Curriculum Committee, Department of Biochemistry and Biophysics, 1993 Chairman, Planning Committee for the UCSF Public Symposium on the DNA Revolution in
Biology, 1992 Executive Committee, UCSF Transgenic Facility, 1993 PUBLICATIONS
Books 1. Varmus, H.E. and Levine, A.J. (eds.) Selected Readings in Tumor Virology . Cold Spring
Harbor Laboratory, 1983. 2. Weiss, R., Teich, N., Varmus, H. and Coffin, J. (eds.) The Molecular Biology of RNA
Tumor Viruses. 1982, 1984.
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3. Varmus, H. and Weinberg, R. A. Genes and the Biology of Cancer. Scientific American
Library, 1992. 4. Stamatoyannopoulos, G., Neinhuis, A.W., Majerus, P.W., Varmus, H.E., (eds.) The
Molecular Basis of Blood Diseases, (2nd edition), 1993. 5. Stamatoyannopoulos, G., Majerus, P.W., Perlmutter, R.M., Varmus, H.E., (eds.) The
Molecular Basis of Blood Diseases, (3rd edition), 2001. 6. Coffin, J., Hughes, S., and Varmus, H., (eds.) Retroviruses. Cold Spring Harbor Laboratory
Press, 1997. 7. Varmus, H. The Art and Politics of Science, W.W. Norton, 2009. Policy (Articles, Letters, Editorials, Lectures, and Chapters) 1. Bishop, J.M., Kirschner, M., Varmus, H.E. Policy Forum: Science and the New
Administration. Science 259:444-445, 1993. 2. Varmus, H.E. Basic Science and the NIH. Molecular Biology of the Cell 5:267-272, 1994. 3. Varmus, H. An overview of the FY-1996 budget for the National Institutes of Health.
Acad. Med. 70:385-387, 1995. 4. Varmus, H. Shattuck Lecture B Biomedical Research Enters the Steady State. N. Engl. J.
Med. 33:811-815, 1995. 5. Varmus, H. An active National Institutes of Mental Health. Science 268:1681, 1995. 6. Varmus, H. NIH review of gene therapy protocols. Science 267:1889, 1995. 7. Varmus, H. and Paul, W.E. Planning of US AIDS research. Nature 381:270, 1996. 8. Varmus, H.E. The Pastorian: A legacy of Louis Pasteur. In Advances in Cancer Research
(G. F. Vande Woude, G. Klein, eds.) Academic Press, pp. 1-16, 1996. 9. Bruno, J.M., Feachem, R., Godal, T., Nchinda, T., Ogilive, B., Mons, B., Mshana, R.,
Radda, G., Samda, E. Schwartz, M., Varmus, H., Diallo, S. Doumbo, O., Greenwood, B., Kilama, W., Miller, L.H., and da Silva, L.P. The spirit of Dakar: a call for action on malaria. Nature 386:541, 1997.
10. Varmus, H. US support for malaria research. Nature 388:416, 1997. 11. Varmus, H. and Satcher, D. Ethical complexities of conducting research in developing
countries. N.Engl. J. Med. 337:1003-1005, 1997.
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12. Varmus, H. and Paul, W. AIDS: the process of discovery. Science 275:141, 1997. 13. Varmus, H. The view from the National Institutes of Health. In The Future of Biomedical
Research (C.E. Barfield and B.L.R. Smith, eds.) The AEI Press, Publisher for the American Enterprise Institute, 1997.
14. Gallin, J.I., Varmus, H. Revitalization of the Warren G. Magnuson Clinical Center at the
National Institutes of Health. Acad Med. 73:460-466, 1998. 15. Varmus, H., Nathanson, N. Science and the Control of AIDS. Science 280:1815, 1998. 16. Varmus, H. National Institutes of Health in the tropics. Am J Trop Med Hyg 59:24-28,
1998. 17. Varmus, H. Evaluating the burden of disease and spending the research dollars of the
National Institutes of Health [editorial; comment]. N. Engl. J. Med. 340:1914-1915, 1998. 18. Nathan, D.G. and Varmus, H.E. The National Institutes of Health and clinical research: a
progress report. Nature Med 6: 1201-1204, 2000.
19. Varmus, H. Policy Forum: Proliferation of National Institutes of Health. Science 291:1903-1905, 2001.
20. Varmus, H.E. The challenge of making laws on the shifting terrain of science. J Law Med
Ethics Suppl.to 28. no. 4:46-53, 2000. 21. Varmus, H. How popular TV shows shape public perceptions of medical scientists. Mt
Sinai J Med 68:346-349, 2001. 22. Varmus, H. Genomic empowerment: the importance of public databases. Nature Gen 32:3,
2002. 23. Varmus, H. Building a global culture of science. The Lancet (Suppl) Medicine and Conflict,
s1-s4, 2002. 24. Varmus, H. What does knowing about genomes mean for science and soceity? In The
Genomic Revolution: Unveiling the Unity of Life (M. Yudell and R. DeSalle, eds.) Joseph Henry Press with the American Museum of Natural History, pp. 20-34, 2002.
25. Varmus, H. Getting ready for gene-based medicine N. Engl. J. Med. 347:1526-1527, 2002. 26. Varmus, H. Book review of The Demon in the Freezer by Richard Preston. The New York
Times Book Review, Vol. CVII, No. 45, pp. 12-13, November 10, 2002. 27. Klausner, R.D., Fauci, A.S., Corey, L., Nabel, G.J., Gayle, H., Berkley, S., Haynes, B.F.,
Baltimore, D., Collins, C., Douglas, R.G., Esparza, J., Francis, D.P., Ganguly, N.K., Gerberding, J.L., Johnston, M.I., Kazatchkine, M.D., McMichael, A.J., Makgoba, M.W.,
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Pantaleo, G., Piot, P., Shao, Y., Tramont, E., Varmus, H., and Wasserheit, J.N. Policy Forum: The need for a global HIV vaccine enterprise. Science 300:2036-2039, 2003.
28. Varmus, H. Film review of 28 Days Later. The New York Times, Section 2, p. 9, July 6,
2003. 29. Brown, P.O., Eisen, M.B., Varmus, H.E. Why PLoS became a publisher. PLoS Biology
2003 October; 1(1):e36. Published online 2003 October 13. 30. Varmus, H., Klausner, R., Zerhouni, E., Acharya, T., Daar, A.S., Singer, P.A. Policy Forum:
Grand Challenges in Global Health. Science 302:398-399, 2003. 31. Varmus, H. Genes and Cancer. DNA: 50 Years of the Double Helix. Cambridge Conference
Extracts. MRC Laboratory of Molecular Biology and Amersham Biosciences, pp.40-46, 2003.
32. Varmus, H. Film review of Godsend. The New York Times, Section 2, p. 18, May 2, 2004. 33. Varmus, H. Television review of Medical Investigation. The New York Times, Section 2, p.
17, September 5, 2004. 34. Bishop, J.M. and Varmus, H. Re-Aim Blame for NIH’s Hard Times. Science 312:499, 2006. 35. Varmus, H. The New Era in Cancer Research. Science 312:1162-1165, 2006.
36. Varmus, H. Time for a Global Science Corps. The Scientist 20:24, 2006. 37. Varmus, H. Global Science Corps on the March. TWAS Newsletter 18 (1) 43-47 (2006). 38. Varmus, H. Is Science Under Siege? AAAS Bulletin Vol. LIX, No. 4, pp. 6-11, Summer
2006. 39. Varmus, H. An Interview with Harold Varmus. Nature Reviews/Molecular Cell Biology 9:502-503, 2008. 40. Gottfried, H., Varmus, H. The Enlightenment Returns. Science 323: 1538, 2009. 41. Varmus, H. and Trimble, E.L. Integrating Cancer Control into Global Health. Science Translational Medicine 3: 34-36, 2011. 42. Cohen, J.I., Fauci, A.S., Varmus, H., Nabel, G.J. Epstein-Barr Virus: An Important Vaccine
Target for Cancer Prevention. Science Translational Medicine 3:107fs7, 2011. 43. Varmus, H. and Harlow E. Provocative Questions in Cancer Research. Nature 481:436-437,
2012. 44. Varmus, H. Arlen Specter: A retrospective. PNAS 109:21178-21179, 2012.
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45. Sacco, R.L., Smith, S.C., Holmes, D., Shurin, S., Brawley, O., Cazap, E., Glass, R., Komajda,
M., Koroshetz, W., Mayer-Davis, E., Mbanya, J.C., Sledge, G., Varmus, H. Accelerating Progress on Non-communicable Diseases. Lancet 382: e4-5, 2013.
46. Varmus, H. and Kumar H.S. Addressing the Growing International Challenge of Cancer: A
Multinational Perspective. Science Translational Medicine 5:175cm2, 2013. 47. Varmus, H. Making PEPFAR: A Triumph of Medical Diplomacy. Science & Diplomacy
2(4) December, 2013. 48. Varmus, H. Medical Research Centers in Mali and Uganda: Overcoming Obstacles to
Building Scientific Capacity and Promoting Global Health. Science & Diplomacy. March, 2014.
49. Alberts, B., Kirschner, M.W., Tilghman, S., Varmus, H. Rescuing U.S. Biomedical Research
from its Systemic Flaws. PNAS 111:5773-5777, 2014. 50. Abrams, J., Kramer, B., Doroshow, J.H., Varmus, H. National Cancer Institute—Supported
Clinical Trials Networks. J. Clin. Oncology 33: 293, 2015. 51. Collins, F.S. and Varmus, H. A New Initiative on Precision Medicine. NEJM 372:793-795,
2015. 52. Varmus, H. Can cancer be stopped? New York Daily News, April 12, 2015.
Science (Articles and Chapters) 1. Bertcher, R.W., Meyer, L.M., Varmus, H.E. and Mulzac, C. Some characteristics of binding
of vitamin B12 by normal human serum. Acta Haematol. 23:287, 1960. 2. deCrombrugghe, B., Perlman, R.L., Varmus, H.E. and Pastan, I. Regulation of inducible
enzyme synthesis in Escherichia coli by cyclic adenosine 3', 5'-monophosphate. J. Biol.Chem.244:5828, 1969.
3. deCrombrugghe, B., Varmus, H.E., Perlman, R.L. and Pastan, I. Stimulation of lac mRNA
synthesis by cyclic AMP in cell free extracts of Escherichia coli. Biochem. Biophys. Res. Commun. 38:894, 1970.
4. Varmus, H.E., deCrombrugghe, B., Perlman, R.L. and Pastan, I. Cyclic 3'-5' adenosine
monophosphate regulation of E. coli messenger RNA synthesis in vivo and in vitro. Fed. Proc. 29:601, 1970.
5. Varmus, H.E., Perlman, R.L. and Pastan, I. Regulation of lac messenger ribonucleic acid
synthesis by cyclic adenosine 3'-5' monophosphate and glucose. J. Biol. Chem. 245:2259, 1970.
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6. Varmus, H.E., Perlman, R.L. and Pastan, I. Regulation of lac transcription Escherichia coli by cyclic adenosine 3', 5'- monophosphate: Studies with DNA-RNA hybridization and hybridization competition. J. Biol. Chem. 245:6366, 1970.
7. Perlman, R.L., Chen, B., deCrombrugghe, B., Emmer, M., Gottesman, M., Varmus, H.E. and
Pastan, I. The role of the cyclic AMP receptor protein in the regulation of lac mRNA synthesis. Cold Spring Harbor Symp. Quant. Biol. 35:419, 1970.
8. Pastan, I., Perlman, R.L., deCrombrugghe, B., Emmer, M. and Varmus, H.E. Regulation of
inducible operons in E. coli by cyclic AMP. In Proceedings of Fogarty International Conference on Adenyl Cyclase, U.S. Government Printing Office, 1970.
9. Varmus, H.E., Perlman, R.L. and Pastan, I. Lac transcription in antibiotic-treated E. coli:
Regulation by cyclic AMP and pseudopolar effects of chloramphenicol and puromycin. Nature New Biology 230:41, 1971.
10. deCrombrugghe, B., Chen, B., Gottesman, M., Pastan, I., Varmus, H.E., Emmer, M. and
Perlman, R.L. Regulation of lac mRNA synthesis by cyclic AMP receptor protein and lac repressor in a soluble cell-free system. Nature New Biology 230:37, 1971.
11. Pastan, I., Perlman, R.L., Emmer, M., Varmus, H.E., deCrombrugghe, B., Chen, B. and
Parks, J. Regulation of gene expression in Escherichia coli by cyclic AMP. Recent Progress in Hormone Research, 27 (Proceedings of the 1970 Laurentian Hormone Conference), Academic Press, 1971.
12. Miller, Z., Varmus, H.E., Parks, J., Perlman, R.L. and Pastan, I. Regulation of gal messenger
ribonucleic acid synthesis in Escherichia coli by cyclic adenosine 3', 5'-monophosphate. J. Biol. Chem. 246:2898, 1971.
13. Bishop, J.M., Varmus, H.E., Levinson, W., Faras, A., Garapin, A., Leong, J., McDonnell, J.
and Taylor, J. RNA-directed DNA polymerase and the replication of Rous sarcoma virus. Fed. Proc. 30:1050, 1971.
14. Varmus, H.E., Levinson, W.E. and Bishop, J.M. Extent of transcription by the RNA-
dependent DNA polymerase of Rous sarcoma virus. Nature New Biology 233:19, 1971. 15. Varmus, H.E., Weiss, R.A., Friis, R., Levinson, W.E. and Bishop, J.M. Detection of avian
tumor virus-specific nucleotide sequences in avian DNAs. Proc. Natl. Acad. Sci. USA 69:20, 1972.
16. Levinson, W.E., Varmus, H.E., Garapin, A.C. and Bishop, J.M. Further characterization of
DNA found in Rous sarcoma virus. Science 175:76, 1972. 17. Taylor, J.M., Faras, A., Varmus, H.E., Levinson, W.E. and Bishop, J.M. RNA-dependent
DNA synthesis by the purified DNA polymerase of Rous sarcoma virus: Characterization of the enzymatic product. Biochemistry 11:2343, 1972.
18. Varmus, H.E., Bishop, J.M., Nowinski, R. and Sarkar, N.H. Mammary tumor virus specific
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nucleotide sequences in mouse DNA. Nature New Biology 238:189-91, 1972. 19. Taylor, J.M., Faras, A.J., Varmus, H.E., Goodman, H.M., Levinson, W.E. and Bishop, J.M.
Transcription of RNA by the RNA-directed DNA polymerase of Rous sarcoma virus and DNA polymerase I of E. coli. Biochemistry 12:460, 1973.
20. Varmus, H.E., Vogt, P.K. and Bishop, J.M. Appearance of virus-specific DNA in mammalian
cells following transformation by Rous sarcoma virus. J. Mol. Biol. 74:613, 1973. 21. Bishop, J.M., Faras, A.J., Garapin, A.C., Hansen, C., Jackson, N., Levinson, W.E., Taylor,
J.M. and Varmus, H.E. RNA directed DNA polymerase and the replication of Rous sarcoma virus. In M.D. Anderson Symposium on The Molecular Basis of Neoplasia, Houston, 1972, The Williams and Wilkins Co., Baltimore, Maryland, 1974, pp. 229-57.
22. Bishop, J.M., Faras, A.J., Garapin, A.C., Goodman, H., Levinson, W.E. Stavnezer, J., Taylor,
J.M. and Varmus, H.E. Characteristics of the transcription of RNA by the DNA polymerase of Rous sarcoma virus. In DNA Synthesis in Vitro, The Steenbock Symposium, (R.D. Wells and R.B. Inman, eds.), University Park Press, Baltimore, 1973, pp. 341-359.
23. Garapin, A.C., Varmus, H.E., Faras, A.J., Levinson, W.E. and Bishop, J.M. RNA-directed
DNA synthesis by virions of Rous sarcoma virus: Further characterization of the templates and the extent of their transcription. Virology 52:264, 1973.
24. Bishop, J.M., Jackson, N., Levinson, W.E., Medeiros, E., Quintrell, N. and Varmus, H.E. The
presence and expression of RNA tumor virus genes in normal and infected cells. Clinical Pathologists Symposium in Amer. J. Clin. Path. 60:31, 1973.
25. Varmus, H.E., Hansen, R.C., Medeiros, E., Deng, C.T. and Bishop, J.M. Detection and
characterization of RNA tumor virus-specific nucleotide sequences in cell DNA. In Possible Episomes in Eukaryotes, Proceedings of Fourth Lepetit Colloquium (L.G. Silvestri, ed.) North Holland/American Elsevier Printing Co., Amsterdam, 1973, pp. 50-60.
26. Bishop, J.M., Jackson, N., Quintrell, N. and Varmus, H.E. Transcription of the RNA tumor
virus genes in normal and infected cells. In Possible Episomes in Eukaryotes, Proceedings of Fourth Lepetit Colloquium (L.C. Silvestri, ed.) North Holland/American Elsevier Printing Co., Amsterdam, 1973, pp. 61-73.
27. Varmus, H.E., Quintrell, N., Medeiros, E., Bishop, J.M., Nowinski, R. and Sarkar, N.H.
Transcription of mouse mammary tumor virus genes in tissues from high and low tumor incidence mouse strains. J. Mol. Biol. 79:663, 1973.
28. Varmus, H.E., Vogt, P.K. and Bishop, J.M. Integration of Deoxyribonucleic acid specific for
Rous sarcoma virus after infection of permissive and nonpermissive hosts. Proc. Natl. Acad. Sci. USA 70:3067, 1973.
29. Varmus, H.E., Bishop, J.M. and Vogt, P.K. Synthesis and integration of Rous sarcoma virus-
specific DNA in permissive and non-permissive hosts. In Proceedings of the 1973 ICN-UCLA Symposium on Virus Research (F. Fox and W.J. Robinson, eds.) Academic Press,
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New York, 1973, pp. 373-379. 30. Bishop, J.M., Deng, C.T., Faras, A.J., Goodman, H.M., Levinson, W.E., Taylor, J.M. and
Varmus, H.E. Transcription of the Rous sarcoma virus genome by RNA-directed DNA polymerase. In Proceedings of the 1973 ICN-UCLA Symposium on Virus Research (F. Fox and W.J. Robinson, eds.) Academic Press, New York, 1973, pp. 15-31.
31. Varmus, H.E. Spirochetal Diseases. In Current Diagnosis and Treatment, Lange Medical
Publishers, Los Altos, 1972, 1973, 1974, 1975, 1976, 1977. 32. Haase, A.T. and Varmus, H.E. Demonstration of a DNA provirus in the lytic growth of Visna
virus. Nature New Biology 245:237. 33. Quintrell, N., Varmus, H.E., Bishop, J.M., Nicolson, M.O. and McAllister, R.M.
Homologies among the nucleotide sequences of the genomes of C-type viruses. Virology 58:568, 1974.
34. Taylor, J.M., Varmus, H.E., Faras, A.J., Levinson, W.E. and Bishop, J.M. Evidence for non-
repetitive subunits in the genome of Rous sarcoma virus. J. Mol. Biol. 84:217, 1974. 35. Haase, A.T., Garapin, A.C., Faras, A.J., Varmus, H.E., and Bishop, J.M. Characterization of
the nucleic acid product of the Visna virus RNA dependent DNA polymerase. Virology 57:251, 1974.
36. Bishop, J.M., Quintrell, N., Medeiros, E. and Varmus, H.E. Of bird and mice and men:
Comments on the use of animal models and molecular hybridization in the search for human tumor viruses. Cancer 34:1421, 1974.
37. Deng, C.T., Boettiger, D., MacPherson, I. and Varmus, H.E. Expression of virus-specific
DNA in revertants of Rous sarcoma virus-transformation BHK-21 cells. Virology 62:512, 1974.
38. Varmus, H.E., Guntaka, R.V., Fan, W.J.W., Heasley, S. and Bishop, J.M. Synthesis of viral
DNA in the cytoplasm of duck embryo fibroblasts and in enucleated cells after infection by avian sarcoma virus. Proc. Natl. Acad. Sci. USA 71:3874, 1974.
39. Varmus, H.E., Heasley, S. and Bishop, J.M. Detection of new and additional virus-specific
DNA sequences in chick embryo fibroblasts after infection by Rous sarcoma virus. J. Virol. 14:895, 1974.
40. Varmus, H.E., Guntaka, R.V., Deng, C.T. and Bishop, J.M. Synthesis, structure and function
of Rous sarcoma virus-specific DNA in permissive and non-permissive cells. Cold Spring Harbor Symp. on Quant. Biol. 39:987, 1974.
41. Levy, J.A., Kazan, P.M. and Varmus, H.E. The importance of DNA size for successful
transfection of chicken embryo fibroblasts. Virology 61:297, 1974. 42. Taylor, J.M., Doxy, A., Kan, Y.W., Varmus, H.E., Lie-Injo, L.E., Ganesan, J. and Todd, D.
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Genetic lesion in homozygous A-thalassemia (hydrops fetalis). Nature 251:392, 1974. 43. Varmus, H.E., Stavnezer, J., Medeiros, E. and Bishop, J.M. Detection and characterization of
RNA tumor virus-specific DNA in cells. In Proceedings of the IVth International Symposium on Comparative Leukemia Research, 1973, Leukemogenesis (Y. Ito and R.M. Dutcher, eds.) Univ. of Tokyo Press, Tokyo/Karger, Basel, pp. 451-461, 1975.
44. Bishop, J.M., Deng, C.T., Faras, A.J., Goodman, H.M., Guntaka, R.V., Levinson, W.E.,
Cordell-Stewart, B., Taylor, J.M. and Varmus, H.E. Transcription of the Rous sarcoma virus genome in vitro and in vivo. In Proceedings of the IVth International Symposium on Comparative Leukemia Research, 1973, Leukemogenesis (Y. Ito and R.M. Dutcher, eds.) Univ. of Tokyo Press, Tokyo/Karger, Basel, pp. 517-523, 1975.
45. Bishop, J.M., Deng, C.T., Faras, A.J., Goodman, H.M., Guntaka, R.V., Levinson, W.E.,
Cordell-Stewart, B., Taylor, J.M. and Varmus, H.E. The provirus of Rous sarcoma virus: Synthesis, integration and transcription. In Proceedings of the Symposium on Tumor Virus-Host Cell Interactions , a NATO Advanced Study Institute held in Monte Carlo, 1973 (A. Kolber, ed.) pp. 255-265, 1975.
46. Varmus, H.E., Guntaka, R.V., Deng, C.T., Domenik, C. and Bishop, J.M. Synthesis and
function of avian sarcoma virus-specific DNA in permissive and non-permissive host cells. In Proceedings of XI International Cancer Congress, Florence, Italy, 1974, Vol. 2, pp. 272-276, Except a Medica, Amsterdam, 1975.
47. Guntaka, R.V., Bishop, J.M. and Varmus, H.E. Synthesis and integration of Rous sarcoma
virus proviral DNA in duck embryo fibroblast cells. In Fundamental Aspects of Neoplasia Symposium, held at the Institute of Microbiology Rutgers State University, (A.A. Gottlieb, ed.) Springer-Verlag, New York, pp. 315-323, 1975.
48. Bishop, J.M. and Varmus, H.E. The molecular biology of RNA tumor viruses. In Cancer: A
Comprehensive Treatise (F.F. Becker, ed.) Plenum Press, 1975, Vol. 2, p. 47. 49. Guntaka, R.V., Mahy, B., Bishop, J.M. and Varmus, H.E. Ethidium bromide inhibits the
appearance of closed circular viral DNA and integration of virus-specific DNA in duck cells infected by avian sarcoma virus. Nature 253:507, 1975.
50. Ringold, G., Lasfargues, E., Bishop, J.M. and Varmus, H.E. Production of mouse mammary
tumor virus by cultured cells in the absence and presence of hormones: Assay by molecular hybridization. Virology 65:135, 1975.
51. Kan, W.Y., Dozy, A.M., Varmus, H.E., Taylor, J.M., Lie-Injo, L.E., Holland, J.P., Ganesan, J.
and Todd, D. Deletion of alpha-globin genes in haemoglobin-H disease demonstrates multiple alpha-globin structural loci. Nature 255:255, 1975.
52. Ringold G., Bishop, J.M. and Varmus, H.E. Stimulation of mouse mammary tumor virus
(MMTV) production by glucocorticoids: Studies on the mechanism of action of dexamethasone. In Proceedings of the Fourth ICN-UCLA Symposium (McMann and Fox, eds.) March 1975, pp. 395-403.
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53. Ringold, G., Yamamoto, K.R., Tomkins, G.M., Bishop, J.M. and Varmus, H.E.
Dexamethasone-mediated induction of mouse mammary tumor virus RNA: A system for studying glucocorticoid action. Cell 6:299, 1975.
54. De La Maza, L.M., Faras, A., Varmus, H.E., Vogt, P.K. and Yunis, J.J. Integration of avian
sarcoma virus specific DNA in mammalian chromatin. Exptl. Cell Res. 93:484, 1975. 55. Kan, Y.M., Holland, J.P., Dozy, A.M. and Varmus, H.E. Demonstration of non-functional ß-
globin mRNA in homozygous ß-thalassemia. Proc. Natl. Acad. Sci. USA 72:5140, 1975. 56. Stehelin, D., Guntaka, R.V., Varmus, H.E. and Bishop, J.M. Purification of DNA
complementary to nucleotide sequence required for neoplastic transformation of fibroblasts by avian sarcoma viruses. J. Mol. Biol. 101:349, 1976.
57. Stehelin, D., Varmus, H.E., Bishop, J.M. and Vogt, P.K. DNA related to the transforming
gene(s) of avian sarcoma viruses is present in normal avian cells. Nature 260:170, 1976. 58. Ringold, G.M., Blair, P.B., Bishop, J.M. and Varmus, H.E. Nucleotide sequence homologies
among mouse mammary tumor viruses. Virology 70:550, 1976. 59. Varmus, H.E. and Shank, P.R. Unintegrated viral DNA is synthesized in the cytoplasm of
avian sarcoma virus transformed duck cells by viral DNA polymerase. J. Virol. 18:567, 1976.
60. Varmus, H.E., Heasley, S., Linn, and Wheeler, K. Use of alkaline sucrose gradients in a
zonal rotor to detect integrated and unintegrated avian sarcoma virus-specific DNA in cells. J. Virol. 18:574, 1976.
61. Friedrich, R., Morris, V., Goodman, H.M., Bishop, J.M. and Varmus, H.E. Differences
between genomes of two strains of mouse mammary tumor virus as shown by partial RNA sequence analysis. Virology 72:330, 1976.
62. Stavnezer, E., Ringold, G., Varmus, H.E. and Bishop, J.M. RNA complementary to the
genome of RNA tumor viruses in virions and virus-producing cells. J. Virol. 20:342, 1976. 63. Varmus, H.E., Stehelin, D., Spector, D., Tal. J., Fujita, D., Padgett, T., Roulland-Dussoix, D.,
Kung, H.-J. and Bishop, J.M. Distribution and function of defined regions of avian tumor virus genomes in viruses and uninfected cells. ICN-UCLA Symposia on Molecular and Cellular Biology, Vol. 4 (D. Baltimore, A.S. Huang and C.F. Fox, eds.) Academic Press, New York, pp. 339-358, 1976.
64. Bishop, J.M., Deng, C.T., Mahy, B.M.J., Quintrell, N., Stavnezer, E., and Varmus, H.E.
Synthesis of viral RNA in cells infected by avian sarcoma viruses. ICN-UCLA Symposia on Molecular and Cellular Biology, Vol. 4 (D. Baltimore, A.S. Huang and C.F. Fox, eds.) Academic Press, New York, pp. 1-20, 1976.
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65. Guntaka, R.V., Richards, O.C., Shank, P.R., Kung, H-J., Davidson, N., Fritsch, E., Bishop, J.M. and Varmus, H.E. Covalently closed circular DNA of avian sarcoma virus: Purification from nuclei of infected quail tumor cells and measurement by electron microscopy and gel electrophoresis. J. Mol. Biol. 106:337, 1976.
66. Verma, I.M., Varmus, H.E. and Hunter, E. Characterization of early temperature-sensitive
mutants of avian sarcoma viruses: Biological properties, thermolability of reverse transcriptase in vitro, and synthesis of viral DNA in infected cells. Virology 74:16, 1976.
67. Bishop, J.M., Stehelin, D., Tal, J., Fujita, D., Roulland-Dussoix, D., Padgett, T., Spector, D.
and Varmus, H.E. The transforming gene of avian sarcoma virus. Symposium on the Molecular Biology of the Mammalian Genetic Apparatus, California Institute of Technology, December, 1975 (P. Ts'o, ed.) Elsevier/North-Holland Biomedical Press, pp. 263-274, 1977.
68. Ringold, G., Yamamoto, K., Shank, P.R. and Varmus, H.E. Mouse mammary tumor virus
DNA in infected rat cells: Characterization of unintegrated forms. Cell 10:19, 1977. 69. Ringold, G., Cardiff, R.D., Varmus, H.E. and Yamamoto, K. Infection of cultured rat
hepatoma cells by mouse mammary tumor virus. Cell 10:11, 1977. 70. Deng, C.T., Stehelin, D., Bishop, J.M. and Varmus, H.E. Characteristics of virus-specific
RNA in avian sarcoma virus-transformed BHK-21 cells and revertants. Virology 76:313, 1977.
71. Stehelin, D., Fujita, D., Padgett, T., Varmus, H.E. and Bishop, J.M. Detection and
enumeration of transformation defective strains of avian sarcoma virus with molecular hybridization. Virology 76:675, 1977.
72. Tal, J., Fujita, D.J., Kawai, S., Varmus, H.E. and Bishop, J.M. Purification of DNA
complementary to the env gene of avian sarcoma virus and analysis of relationships among the env genes of avian leukosis-sarcoma viruses. J. Virol. 21:497, 1977.
73. Morris, V., Medeiros, E., Ringold, G., Bishop, J.M. and Varmus, H.E. Comparison of mouse
mammary tumor virus-specific DNA in inbred, wild, and Asian mice and in tumors and normal organs from inbred mice. J. Mol. Biol. 114:7391, 1977.
74. Bister, K., Varmus, H.E., Stavnezer, E., Hunter, E. and Vogt, P.K. Biological and
biochemical studies on the inactivation of avian oncoviruses by ultraviolet irradiation. Virology 77:689, 1977.
75. Varmus, H.E., Spector, D.H., Stehelin, D., Deng, C.T., Padgett, T., Stubblefield, E. and
Bishop, J.M. The function and origin of the transforming gene of avian sarcoma virus. Cold Spring Harbor Conferences on Cell Proliferation, Vol. 4, Cold Spring Harbor Press, pp. 1159-1168, 1977.
76. Padgett, T., Stubblefield, E. and Varmus, H.E. Chicken macrochromosomes contain an
endogenous provirus and microchromosomes contain sequences related to the transforming gene of ASV. Cell 10:649, 1977.
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77. Varmus, H.E., Spector, D.H., Stehelin, D., Deng, C.T., Padgett, T., Stubblefield, E. and
Bishop, J.M. The function and origin of the transforming gene of avian sarcoma virus. International Cell Biology, 1976-1977 (B.R. Brinkley and K.R. Porter, eds.) Rockefeller Univ. Press, pp. 553-7.
78. Tal, J., Kung, H-J., Varmus, H.E. and Bishop, J.M. Characterization of DNA complementary
to nucleotide sequences adjacent to poly(A) at the 3' terminus of the avian sarcoma virus genome. Virology 79:183, 1977.
79. Varmus, H.E., Shank, P.R., Kung, H-J., Guntaka, R.V., Richards, O., and Bishop, J.M.
Mechanism of synthesis and persistence of virus-specific DNA in cells infected by RNA tumor virus. Microbiology-1977 (D. Schlesinger, ed.) pp. 470-2, Am. Soc. Microbiology, 1977.
80. Varmus, H.E., Padgett, T., Heasley, S., Simon, G. and Bishop, J.M. Cellular functions re
required for the synthesis and integration of avian sarcoma virus-specific DNA. Cell 11:307-19, 1977.
81. Weiss, S.R., Varmus, H.E. and Bishop, J.M. The size and genetic composition of virus-
specific RNAs in the cytoplasm of cells producing avian sarcoma-leukosis viruses. Cell 12:983-92, 1977.
82. Oppermann, H., Bishop, J.M., Varmus, H.E. and Levintow, L. A joint product of the genes
gag and pol of avian sarcoma virus: A possible precursor of reverse transcriptase. Cell 12:993-1005, 1977.
83. Ringold, G.M., Yamamoto, K.R., Bishop, J.M. and Varmus, H.E. Glucocorticoid stimulated
accumulation of mouse mammary tumor virus RNA: Increased rate of synthesis of viral RNA. Proc. Natl. Acad. Sci. USA 74:2879, 1977.
84. Friedrich, R., Kung, H-J., Baker, B., Varmus, H.E., Goodman, H.M. and Bishop, J.M.
Characterization of DNA complementary to nucleotide sequences to the 5' terminus of the avian sarcoma virus genome. Virology 79:198, 1977.
85. Spector, D.H., Varmus, H.E. and Bishop, J.M. Nucleotide sequences related to the
transforming gene of avian sarcoma virus are present in the DNA of uninfected vertebrates. Proc. Natl. Acad. Sci. USA 75:4102, 1978.
86. Bishop, J.M., Baker, B., Fujita, D., McCombe, P., Sheiness, D., Smith, K., Spector, D.H.,
Stehelin, D. and Varmus, H.E. The genesis of a viral transforming gene. 3rd Decentennial Review Conference: Cell Tissue, and Organ Culture, held in Lake Placid, New York, September 13-17, 1976. NCI Monograph No. 48, pp. 219-23, 1978.
87. Spector, D., Baker, B., Varmus, H.E. and Bishop, J.M. Characteristics of cellular RNA
related to the transforming gene of avian sarcoma viruses. Cell 3:381, 1978. 88. Spector, D., Smith, K., Padgett, T., McCombe, P., Roulland-Dussoix, D., Moscovici, C.,
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Varmus, H.E. and Bishop, J.M. Uninfected avian cells contain RNA related to the transforming gene of avian sarcoma viruses. Cell 3:371, 1978.
89. Shank, P.R. and Varmus, H.E. Virus-specific DNA in the cytoplasm of avian sarcoma virus
infected cells is a precursor to covalently closed circular viral DNA in the nucleus. J. Virol. 25:104, 1978.
90. Varmus, H.E., Heasley, S., Kung, H.J., Oppermann, H., Smith, V.C., Bishop, J.M. and Shank,
P.R. Kinetics of synthesis, structure, and purification of avian sarcoma virus-specific DNA made in the cytoplasm of acutely-infected cells. J. Mol. Biol. 120:55, 1978.
91. Shank, P.R., Cohen, J.C., Varmus, H.E., Yamamoto, K.R. and Ringold, G.M. Mapping of
linear and circular forms of mouse mammary tumor virus DNA with restriction endonucleases: Evidence for a large specific deletion occurring at high frequency during circularization. Proc. Natl. Acad. Sci. USA 75:2112, 1978.
92. Bishop, J.M., Weiss, S.R., Oppermann, H., Hackett, P., Quintrell, N., Chen, L.S., Levintow,
L. and Varmus, H.E. The strategy of retrovirus gene expression. Avian RNA Tumor Viruses (S. Barlati and C. De Giuli-Morghen, eds.) Piccin Medical Books, pp. 181-189, 1978.
93. Fujita, D., Tal, T., Varmus, H.E. and Bishop, J.M. The env gene of chicken RNA tumor
viruses: Extent of conservation in cellular and viral genomes. J. Virol. 27:465, 1978. 94. Cordell, B., Weiss, S., Varmus, H.E. and Bishop, J.M. A sequences of at least 104
nucleotides in transposed from the 5' terminus of the avian sarcoma virus genome to the 5' termini of smaller viral mRNAs. Cell 15:79, 1978.
95. Levinson, A., Oppermann, H., Levintow, L., Varmus, H.E. and Bishop, J.M. Evidence that
the transforming gene of avian sarcoma virus encodes a protein kinase associated with a phosphoprotein. Cell 15:561, 1978.
96. Shank, P.R., Hughes, S., Kung, H.J., Majors, J., Quintrell, N., Guntaka, R.V., Bishop, J.M.
and Varmus, H.E. Mapping unintegrated avian sarcoma virus DNA; Termini of linear DNA bear 300 nucleotides present once or twice in two species of circular DNA. Cell 15:1383, 1978.
97. Hughes, S., Vogt, P.K., Shank, P.R., Spector, D., Kung, H.J., Breitman, M.L., Bishop, J.M.
and Varmus, H.E. Proviruses of avian sarcoma virus are terminally redundant, co-extensive with unintegrated linear DNA, and integrated at many sites in rat cell DNA. Cell 15:1397, 1978.
98. Varmus, H.E., Shank, P.R., Hughes, S., Kung, H.J., Heasley, S., Majors, J., Vogt, P.K. and
Bishop, J.M. Synthesis, structure and integration of the DNA of RNA tumor viruses. Cold Spring Harbor Symp. Quant. Biol. 43:851, 1979.
99. Varmus, H.E., Cohen, J.C., Shank, P.R., Ringold, G.M., Yamamoto, K.R., Cardiff, R. and
Morris, V. The endogenous and acquired proviruses of mouse mammary tumor virus. ICN-UCLA Symposium (G. Todaro, J. Stevens, and F. Fox, eds.) Academic Press, pp. 161-179,
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1979. 100. Varmus, H.E., Ringold, G. and Yamamoto, K.R. Regulation of mouse mammary tumor virus
gene expression by glucocorticoid hormones. Glucocorticoid Hormone Action (J. Baxter and G. Rousseau, eds.) Springer-Verlag, Vol. 12, pp. 253-278, 1979.
101. Cohen, J.C., Shank, P.R., Morris, V., Cardiff, R. and Varmus, H.E. Integration of DNA from
mouse mammary tumor viruses in normal and neoplastic tissue of the mouse. Cell 16:333, 1979.
102. Varmus, H.E., Shank, P.R., Hughes, S.H., Kung, H.J., Heasley, S., Majors, J., Vogt, P.K. and
Bishop, J.M. Structures of unintegrated and integrated forms of the DNA of RNA tumor viruses. In Symposium of the Biochemical Society, Biochemistry of Genetic Engineering 44:103, 1979.
103. Robertson, D. and Varmus, H.E. Structure and function of the intracellular RNAs of murine
mammary tumor virus. J. Virol. 30:576, 1979. 104. Hughes, S.H., Stubblefield, E., Payvar, F., Engel, J.D., Dodgson, J.B., Spector, D., Cordell,
B., Schimke, R.T. and Varmus, H.E. Gene localization by chromosome fractionation: Globin genes are on at least two chromosomes and three estrogen inducible genes are on three chromosomes. Proc. Natl. Acad. Sci. 76:1348, 1979.
105. Hughes, S.H., Payvar, F., Spector, D., Schimke, R.T., Robinson, H.L., Bishop, J.M. and
Varmus, H.E. Heterogeneity of genetic loci in chickens: Analysis of endogenous viral and non-viral genes by cleavage of DNA with restriction endonucleases. Cell 18:347, 1979.
106. Cohen, J.C. and Varmus, H.E. Endogenous mammary tumor virus DNA varies among wild
mice and segregates during inbreeding. Nature 278:418, 1979. 107. Oppermann, H., Levinson, A.D., Varmus, H.E., Levintow, L. and Bishop, J.M. Uninfected
vertebrate cells contain a protein that is closely related to the product of the avian sarcoma virus transforming gene (src). Proc. Natl. Acad. Sci. USA 76:1804, 1979.
108. Morris, V.L., Kozak, C., Cohen, J.C., Shank, P.R., Jolicoeur, J.,Ruddle, F. and Varmus, H.E.
Endogenous mouse mammary tumor virus DNA is distributed among multiple mouse chromosomes. Virology 92:46, 1979.
109. Ringold, G.M., Shank, P.R., Varmus, H.E., Ring, J. and Yamamoto, K.R. Integration and
transcription of mouse mammary tumor virus DNA in rat hepatoma cells. Proc. Natl. Acad. Sci. USA 76:665, 1979.
110. Lee, J.S., Varmus, H.E. and Bishop, J.M. Virus-specific messenger RNAs in permissive cells
infected by avian sarcoma virus. J. Biol. Chem. 254:8015, 1979. 111. Martin, G.S., Radke, K., Hughes, S., Quintrell, N., Bishop, J.M. and Varmus, H.E. Mutants
of Rous sarcoma virus containing extensive deletions of the viral genome. Virology 96:530, 1979.
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112. Hughes, S.H., Robinson, H.L., Bishop, J.M. and Varmus, H.E. The replication of subgroup E
avian retroviruses is blocked at or before viral DNA synthesis in restrictive chicken cells. Virology 99:437, 1979.
113. Cohen, J.C., Majors, J.E. and Varmus, H.E. Organization of mouse tumor virus-specific
DNA endogenous to BALB/c mice. J. Virol. 32:483, 1979. 114. Hughes, S.H., Vogt, P.K., Stubblefield, E., Robinson, H., Bishop, J.M. and Varmus, H.E.
Organization of endogenous and exogenous viral and linked non-viral sequences. Cold Spring Harbor Symp. Quant. Biol. 44:1077, 1980.
115. Bishop, J.M., Courtneidge, S.A., Levinson, A.R., Oppermann, H., Quintrell, N., Sheiness,
D.K., Weiss, S.R. and Varmus, H.E. The origin and function of avian retrovirus transforming genes. Cold Spring Harbor Symp. Quant. Biol. 44:919, 1980.
116. Hughes, S.H., Stubblefield, E., Nazarian, K. and Varmus, H.E. DNA of a chicken herpesvirus
is associated with at least two chromosomes in a chicken lymphoblastoid cell line. Virology 105:234, 1980.
117. Varmus, H.E., Cohen, J.C., Hughes, S.H., Majors, J. and Bishop, J.M. The origin and
structure of endogenous retroviral DNA. Annals of New York Academy of Sciences - Genetic Variation of Viruses (Jan. 1980), Vol. 354, pp. 379-383, 1980.
118. Sheiness, D.K., Hughes, S.H., Varmus, H.E., Stubblefield, E. and Bishop, J.M. The
vertebrate homologue of the putative transforming gene of avian myelocytomatosis virus: Characteristics of the DNA locus and its RNA transcript. Virology 105:415, 1980.
119. Bishop, J.M., Gonda, T., Hughes, S.H., Sheiness, D.K., Stubblefield, E. Vennstrom, B. and
Varmus, H.E. The genesis of avian retrovirus oncogenes. Mobilization and Reassembly of Genetic Information (Twelfth Miami Winter Symp.) January, 1980, pp. 261-273.
120. Quintrell, N., Hughes, S.H., Varmus, H.E. and Bishop, J.M. The structure of viral DNA and
RNA in mammalian cells infected with avian sarcoma virus. J. Mol. Biol. 143:363, 1980. 121. Cohen, J.C. and Varmus, H.E. Proviruses of mouse mammary tumor virus in normal and
neoplastic tissues from GR and C3Hf mouse strains. J. Virol. 35:298, 1980. 122. Czernilofsky, A.P., DeLorbe, W., Swanstrom, R., Varmus, H.E., Bishop, J.M., Tischer, E. and
Goodman, H.M. The nucleotide sequence of an untranslated but conserved domain at the 3' end of the avian sarcoma virus. Nucl. Acids Res. 8:2967, 1980.
123. Kung, H.J., Shank, P.R., Bishop, J.M. and Varmus, H.E. Identification and characterization
of dimeric and trimeric circular forms of avian sarcoma virus-specific DNA. Virology 103:425, 1980.
124. Majors, J.E. and Varmus, H.E. Learning about the replication of retroviruses from a single
cloned provirus of mouse mammary tumor virus. ICN-UCLA Symposium (B. Field, R.
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Jaenisch and C.F. Fox, eds.) Academic Press, pp. 241-253, 1980. 125. Wyke, J.A., Beamand, J.A. and Varmus, H.E. Factors affecting phenotypic reversion of rat
cells transformed by avian sarcoma virus. Cold Spring Harbor Symp. Quant. Biol. 44:1065, 1980.
126. DeLorbe, W.J., Luciw, P.A., Varmus, H.E., Bishop, J.M. and Goodman, H.M. Molecular
cloning and characterization of avian sarcoma virus circular DNA molecules. J. Virol., 36:50, 1980.
127. Levinson, A.D., Oppermann, H., Varmus, H.E. and Bishop, J.M. The purified product of the
transforming gene of avian sarcoma virus phosphorylates tyrosine. J. Biol. Chem. 255:11973, 1980.
128. Czernilofsky, A.P., Levinson, A.D., Varmus, H.E., Bishop, J.M., Tischer, E. and Goodman,
H.M. The oncogene of avian sarcoma virus (src): Nucleotide sequence of the gene and a proposed amino acid sequence for the gene product (pp60ÆMDSUØsrc). Nature 287:198, 1980.
129. Majors, J.E., Swanstrom, R., DeLorbe, W.J., Payne, G.S., Hughes, S.H., Ortiz, S., Quintrell,
N., Bishop, J.M. and Varmus. H.E. DNA intermediates in the replication of retroviruses are structurally (and perhaps functionally) related to transposable elements. Cold Spring Harbor Symp. Quant. Biol. 45:731, 1980.
130. Hughes, H.S., Toyoshima, K., Bishop, J.M. and Varmus, H.E. Organization of the
endogenous proviruses of chickens: Implications for origin and expression. Virology 108:189, 1981.
131. Hughes, S.H., Vogt, P.K., Stubblefield, E., Bishop, J.M. and Varmus, H.E. Integration of
avian sarcoma virus DNA in chicken cells. Virology 108:208, 1981. 132. Shank, P.R., Hughes, S.H. and Varmus, H.E. Restriction endonuclease mapping of the DNA
of Rous-associated virus-0 reveals extensive homology in structure and sequence with avian sarcoma virus DNA. Virology 108:177, 1981.
133. Hughes, S.H., Vogt, P.K., Bishop, J.M. and Varmus, H.E. Endogenous proviruses of random
bred chickens and ring-necked pheasants: Analysis with restriction endonucleases. Virology 108:222, 1981.
134. Varmus, H.E., Quintrell, N. and Wyke, J. Revertants of an ASV-transformed rat cell line
have lost the complete provirus or sustained mutations in src. Virology 108:28, 1981. 135. Bishop, J.M., Courtneidge, S., Czernilofsky, A.P., Levinson, A.D., Levintow, L., Luciw, P.,
Oppermann, H., Sheiness, D.K., Vennstrom, B. and Varmus, H.E. Viral oncogenes as pleiotropic effectors. Genes, Chromosomes, and Neoplasia (33rd Ann. Symp. on Fundamental Cancer Res., M.D. Anderson Series) (F.E. Arrighi, P.N. Rao and E. Stubblefield, eds.) Raven Press, New York, pp. 93-103, 1981.
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136. Swanstrom, R., Varmus, H.E. and Bishop, J.M. The terminal redundancy of the retrovirus genome facilitates chain elongation by reverse transcriptase. J. Biol. Chem. 78:124, 1981.
137. Oppermann, H., Levinson, A.D. and Varmus, H.E. The structure and protein kinase activity
of proteins encoded by non-conditional mutants and back mutants in the src gene of avian sarcoma virus. Virology 108:47, 1981.
138. Kung, H.J., Fung, Y.K., Majors, J.E., Bishop, J.M. and Varmus, H.E. Synthesis of plus
strands of retroviral DNA in cells infected with avian sarcoma virus and mouse mammary tumor virus. J. Virol. 37:127, 1981.
139. Varmus, H.E., Majors, J.E., Swanstrom, R., DeLorbe, W.J., Payne, G.S., Hughes, S.H., Ortiz,
S., Quintrell, N. and Bishop, J.M. Structural and functional properties of DNA intermediates in the replication of retroviruses. Juselius Symposium, Academic Press, pp. 93-109, 1981.
140. Majors, J.E. and Varmus, H.E. Nucleotide sequences at host-proviral junctions for mouse
mammary tumour virus. Nature 289:253, 1981. 141. Swanstrom, R., DeLorbe, W.J., Bishop, J.M. and Varmus, H.E. Sequencing of cloned
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termination of transcription in chicken embryo cells irradiated with ultraviolet light. Virology 112:752, 1981.
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H.E. Analyses of avian leukosis virus DNA and RNA in bursal tumors: viral gene expression is not required for maintenance of the tumor state. Cell 23:311, 1981.
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148. Dudley, J.P. and Varmus, H.E. Purification and translation of murine mammary tumor virus
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retroviruses DNA and RNA: Viral and cellular determinants of retrovirus gene expression. Virology 114:8, 1981.
150. Varmus, H.E., Quintrell, N. and Ortiz, S. Retroviruses as mutagens: Insertion and excision of
a non-transforming provirus alters expression of a resident transforming provirus. Cell 25:23, 1981.
151. Robertson, D.L. and Varmus, H.E. Dexamethasone induction of the intracellular RNAs and
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is flanked by short direct repeats of cellular DNA which were present in only one copy prior to integration. Proc. Natl. Acad. Sci. USA 78:4299, 1981.
153. Parker, R.C., Varmus, H.E. and Bishop, J.M. The cellular homologue (c-src) of the
transforming gene of Rous sarcoma virus: Isolation, mapping, and transcriptional analysis of c-src and flanking regions. Proc. Natl. Acad. Sci. USA 78:5842, 1981.
154. Coffin, J.M., Varmus, H.E., Bishop, J.M., Essex, M., Hardy, Jr., W.D., Martin, G.S.,
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155. Hackett, P.B., Swanstrom, R., Varmus, H.E. and Bishop, J.M. The leader sequence of the
subgenomic messenger RNAs of Rous sarcoma virus is approximately 390 nucleotides. J. Virol. 41:527, 1982.
156. Swanstrom, R., Varmus, H.E. and Bishop, J.M. Nucleotide sequence of the 5' noncoding
region and part of the gag gene of Rous sarcoma virus. J. Virol. 41:535, 1982. 157. Payne, G.S., Bishop, J.M. and Varmus, H.E. Multiple arrangements of viral DNA and an
activated host oncogene (c-myc) in bursal lymphomas. Nature 295:209, 1982. 158. Varmus, H.E. Form and function of retroviral proviruses. Science 216:812, 1982. 159. Swanstrom, R., Bishop, J.M. and Varmus, H.E. Structure of a replication intermediate in the
synthesis of Rous sarcoma virus DNA in vivo. J. Virol. 42:337. 1982. 160. Varmus, H.E. How long terminal repeats in proviral DNA affect the behavior of retroviruses.
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162. Ganem, D., Greenbaum, L., and Varmus, H.E. Virion DNA of ground squirrel hepatitis virus:
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contain a provirus integrated in the same region of the host genome. Cell 31:99, 1982. 166. Varmus, H.E. and Swanstrom, R. Replication of retroviruses. Chapter 5 in Molecular Biology
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suggests src contains two functional sites for the initiation of protein synthesis from a single mRNA. Cell 32:871, 1983.
172. Luciw, P., Bishop, J.M., Varmus, H.E. and Capecchi, M. Location and function of retroviral
and SV40 sequences that enhance biochemical transformation after microinjection of DNA. Cell 33:705, 1983.
173. Alitalo, K., Schwab, M., Lin, C.C., Varmus, H.E. and Bishop, J.M. Homogeneously staining
chromosomal regions contain amplified copies of an abundantly cellular oncogene (c-myc) in malignant neuroendocrine cells from a human colon carcinoma. Proc. Natl. Acad. Sci. USA 80:1707, 1983.
174. Swanstrom, R., Parker, R.C., Varmus, H.E. and Bishop, J.M. Transduction of a cellular
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amplified, overexpressed, and located within karyotypic abnormalities in mouse
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mRNA defines determinants of expression of the MMTV env gene. J. Virol. 47:495, 1983. 177. Weiser, B., Ganem, D., Seeger, C. and Varmus, H.E. Closed circular viral DNA and
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178. Varmus, H.E. and Levine, A.J. (eds.) Selected Readings in Tumor Virology . Cold Spring
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terminal repeat confers glucocorticoid hormone regulation on a linked heterologous gene. Proc. Natl. Acad. Sci. USA 80:5866, 1983.
180. Schwab, M., Alitalo, K., Klempnauer, K.-H., Varmus, H.E., Bishop, J.M., Gilbert, F.,
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181. Nusse, R., van Ooyen, A., Cox, D., Fung, Y.W. and Varmus, H.E. Mode of proviral
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183. Westaway, D., Payne, G., and Varmus, H.E. Proviral deletions and oncogene base-
substitutions in insertionally mutagenized c-myc alleles may contribute to the progression of avian bursal tumors. Proc. Natl. Acad. Sci. USA 81:843, 1984.
184. Brodeur, G.M., Seeger, R.C., Schwab, M., Varmus, H.E., and Bishop, J.M. Amplification of
N-myc in untreated human neuroblastomas correlates with advanced disease stage. Science 224:1121, 1984.
185. Luciw, P., Oppermann, H., Bishop, J.M., and Varmus, H.E. Integration and expression of
several molecular forms of Rous sarcoma virus DNA used for transfection of mouse cells. Mol. Cell. Biol. 4:1260, 1984.
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surface antigen gene in cultured murine cells initiates within the presurface region. J. Virol. 50:563-571, 1984.
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codon in pol is defective for integration. Proc. Natl. Acad. Sci. USA 81:6461, 1984. 194. Parker, R.P., Varmus, H.E., and Bishop, J.M. Expression of v-src and chicken c-src in rat
cells demonstrates qualitative differences between pp60v-src and pp60c-src. Cell 37:131, 1984.
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expression of the human gene N-myc consequent to amplification of DNA may contribute to malignant progression of neuroblastoma. Proc. Natl. Acad. Sci. USA, 81:4940, 1984.
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199. Parker, R.C., Mardon, G., Lebo, R.V., Varmus, H.E., and Bishop, J.M. Isolation of
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200. Mossie, K.G., Young, M.W., and Varmus, H.E. Presence and variation of extrachromosomal
DNA forms of copia-like transposable elements, F-elements, and middle repetitive sequences in cultured cells and embryos of Drosophila melanogaster. J. Mol. Biol. 182:31, 1985.
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204. Persing, D.H., Varmus, H.E. and Ganem, D. A frameshift mutation in the pre-S region of the
human hepatitis B virus genome allows production of surface antigen particles but eliminates binding to polymerized albumin. Proc. Natl. Acad. Sci. USA 82:3440-4, 1985.
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209. Lee, W.M.F., Schwab, M., Westaway, D., and Varmus, H.E. Augmented expression of
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210. Fung, Y.-K.T., Shackleford, G.M., Brown, A.M.C., Sanders, G.S., and Varmus, H.E.
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266. Varmus, H.E. Retroviruses and oncogenes - I. In Les Prix Nobel, 1990 and In Angew.
Chem., 29:707-15, 1990. 267. Hirai, H., Varmus, H.E. SH2 and SH3 mutants of activated chicken C-src that preferentially
transform mouse or chicken cells. Proc. Natl. Acad. Sci. 87:8592-6, 1990. 268. Young, J.A.T., Bates, P., Willert, K., Varmus, H.E. Efficient incorporation of human CD4
protein into avian leukosis virus particles. Science 250:1421-3, 1990. 269. Hirai, H., Varmus, H.E. SH2 mutants of c-src that are host-dependent for transformation are
trans-dominant inhibitors of mouse cell transformation by activated c-src. Genes and Development 4:2342-52, 1990.
270. Varmus, H.E., Retroviruses and oncogenes. Biosci Rep., 10:413-30, 1990. 271. Morgan, D.O., Kaplan, J.M., Bishop, M.J. and Varmus, H.E. Production of p60c-src by
baculovirus expression and immunoaffinity purification. Methods in Enzymology, 200:645-60, 1991.
272. Seeger, C., Baldwin, B., Hornbuckle, W.F., Yeager, A., Tennant, B., Cote, P., Ferrell, L.,
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273. Chamorro, M., Parkin, N., Varmus, H.E. An RNA pseudoknot and an optimal heptameric
shift site are required for highly efficient ribosomal frameshifting on a retroviral messenger RNA. Proc. Natl. Acad. Sci., USA 89:713-7, 1992.
274. Kwan, H., Pecenka, V., Tsukamoto, A., Parslow, T.G., Guzman, R., Lin, T-P., Muller, W.J.,
Lee, F.S., Leder, P., Varmus, H.E. Transgenes expressing the Wnt-1 and int-2 proto-oncogenes cooperate during mammary carcinogenesis in doubly transgenic mice. Mol. and Cell. Biol. 12:147-54, 1992.
275. Jue, S.F., Bradley, R.S., Rudnicki, J.A., Varmus, H.E., Brown, A.M.C. The mouse Wnt-1
gene can act via a paracrine mechanism in transformation of mammary epithelial cells. Mol. Cell. Biol. 12:321-8, 1992.
276. Kitajewski, J., Mason, J.O., Varmus, H.E. Interaction of Wnt-1 proteins with binding protein,
BiP. Mol. Cell. Biol. 12:784-90, 1992. 277. Pryciak, P.M., Sil, A., Varmus, H.E. Retroviral integration into minichromosomes in vitro.
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278. Bourne, H.R. Varmus, H.E. Oncogenes and cell proliferation. Current Opinion in Genetics and Development 2:1-3, 1992.
279. Leavitt, A.D., Rose, R.B., Varmus, H.E. Both substrate and target oligonucleotide sequences
affect in vitro integration mediated by HIV-1 integrase protein (IN) produced in Saccharomyces cerevisiae. J.Virology 66:2359-68, 1992.
280. Pryciak, P. and Varmus, H.E. Nucleosomes, DNA-binding proteins, and DNA sequence
modulate retroviral integration target site selection. Cell 69:769-80, 1992. 281. Parkin, N.T., Chamorro, M., Varmus, H.E. Human immunodeficiency virus type 1 gag-pol
frameshifting is dependent on downstream messenger RNA secondary structure: Demonstration by expression in vivo. J. Virology 66(8):5147-51, 1992.
282. Mason, J.O., Kitajewski, J., Varmus, H.E. Mutational analysis of mouse Wnt-1 identifies two
temperature-sensitive alleles and attributes of Wnt-1 protein essential for transformation of a mammary cell line. Molecular Biology of the Cell 3, 521-33, 1992.
283. Nusse, R., Varmus, H.E. Wnt genes. Cell 69:1073-88, 1992. 284. Pryciak, P., Muller, H.-P., Varmus, H.E. SV40 minichromosomes as targets for retroviral
integration in vivo. Proc. Natl. Acad. Sci., USA, 89:9237-41, 1992. 285. Pryciak, P. and Varmus, H.E. FV-1 restriction and its effects on murine leukemia virus
integration in vivo and in vitro. J. Virol. 66:5959-66, 1992. 286. Kaplan, K.B., Swedlow, J.R., Varmus, H.E., Morgan, D.O. Association of p60c-src with
endosomal membranes in mammalian fibroblasts. J. Cell Biol. 118:321-33, 1992. 287. Leavitt, A.D., Shiue, L., Varmus, H.E. Site-directed mutagenesis of HIV-1 integrase
demonstrates that integrase functions are separable in vitro. J. Biol. Chem. 268:2113-9, 1993.
288. Shackleford, G.M., Willert, K., Wang, J., Varmus, H.E. Wnt-1 proto-oncogene induces
changes in morphology, gene expression and NGF responsiveness, in PC12 cells. Neuron 11:865-75, 1993.
289. Young, J.A.T., Bates, P., Varmus, H.E. Isolation of the chicken gene encoding the receptor
for subgroup A-specific avian leukosis and sarcoma virus. J. Virol. 67:1811-6, 1993. 290. Shackleford, G., Shivakumar, S., Shiue, L., Mason, J., Kenyon, C., Varmus, H.E. cDNA
sequences and developmental expression of two Wnt genes in Caenorhabditis elegans. Oncogene 8:1857-64, 1993.
291. Shackleford, G.M., MacArthur, C.A., Kwan, H.C., Varmus, H.E. Mouse mammary tumor
virus infection accelerates mammary carcinogenesis in Wnt-1 transgenic mice by insertional activation of int-2/Fgf-3 and hst//Fgf-4. Proc. Natl. Acad. Sci. 90:740-4, 1993.
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292. Bates, P., Young, J.A.T., Varmus, H.E. A putative receptor for subgroup A Rous sarcoma
virus is related to the LDL receptor. Cell 74:1043-51, 1993. 293. Bibbins, K.B., Boeuf, H., and Varmus, H.E. Binding of the src SH2 domain to
phosphopeptides is determined by residues in both the SH2 domain and the phosphopeptides. Mol. Cell. Biol. 13:7278-87, 1993.
294. Muller, H-P., Pryciak, P.M., Varmus, H.E. Retroviral integration machinery as a probe for
DNA structure and associated proteins. Cold Spring Harbor Symposia on Quantitative Biology, 533-41, 1993.
295. Parkin, N.T., Kitajewski, J., Varmus, H.E. Activity of Wnt-1 as a transmembrane protein.
Genes & Development 7:2181-93, 1993. 296. Lowell, C.A., Soriano, P., Varmus, H.E. Functional overlap in the src gene family:
inactivation of hck and fgr impairs natural immunity. Genes & Development 8:387-98, 1994. 297. Verderame, M.F. and Varmus, H.E. Highly conserved amino acids in the SH2 and catalytic
domains of v-src are altered in naturally occurring, transformation-defective alleles. Oncogene 9:175-82, 1994
298. Varmus, H.E. and Lowell, C.A. Cancer genes and hematopoiesis. Blood 83:5-9, 1994. 299. Federspiel, M.J., Bates, P., Young, J.A.T., Varmus, H.E., Hughes, S.H. A system for tissue-
specific gene targeting: Transgenic mice susceptible to subgroup A avian leukosis virus-based retroviral vectors. Proc. Natl. Acad. Sci. 91:11241-5, 1994
300. Müller, H-P., Varmus, H.E. DNA bending creates favored sites for retroviral integration: an
explanation for preferred insertion sites in nucleosomes. EMBO J. 13:4704-14, 1994. 301. Kaplan, K.B., Bibbins, K.B., Swedlow, J.R., Arnaud, M., Morgan, D.O., Varmus, H.E.
Association of the amino-terminal half of c-Src with focal adhesions alters their properties and is regulated by phosphorylation of tyrosine 527. EMBO J. 13:4745-56, 1994.
302. Gilbert, J.M., Bates, P., Varmus, H.E., White, J.M. The receptor for the subgroup A avian
leukosis-sarcoma viruses binds to subgroup A but not to subgroup C envelope glycoprotein. J Virol. 68:5623-8, 1994.
303. Varmus, H.E., Godley, L.A., Roy, S., Taylor, I.C.A., Yuschenkoff, L., Shi, Y.P., Pinkel, D.,
Gray, J., Pyle, R., Aldaz, C.M. Bradley, A., Medina, D., Donehower, L.A. Defining the steps in a multistep mouse model for mammary carcinogenesis. Cold Spring Harbor Symposium on Quantitative Biology, 59:491-9, 1994.
304. Lee, S.I., Umen, J., Varmus, H.E. A genetic screen identifies cellular factors involved in
retroviral -1 frameshifting. Proc. Natl. Acad. Sci. 92:6587-91, 1995. 305. Boeuf, H., Murphy, J., Bibbins, K., Varmus, H.E. Binding of phosphotyrosine-containing
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proteins to pp60c-src SH2 domain does not necessarily correlate with cell transformation. Oncogene 10:433-8, 1995
306. Chen, X., Chamorro, M., Lee, S.I., Shen, L.X., Hines, J.V., Tinoco, Jr., I., Varmus, H.E.
Structural and functional studies of retroviral RNA pseudoknots involved in ribosomal frameshifting: nucleotides at the junction of the two stems are important for efficient ribosomal frameshifting. EMBO J. 14:842-52, 1995
307. He X., Saint-Jeannet, J.P., Woodgett, J.R., Varmus, H.E., Dawid, I.B. Glycogen synthase
kinase-3 and dorsoventral patterning in Xenopus embryos. Nature 374:617-22, 1995. 308. Donehower, L.A., Godley, L.A., Aldaz, C.M., Pyle, R., Shi, Y.P., Pinkel, D., Gray, J.,
Bradley, A., Medina, D., Varmus, H.E. Deficiency of p53 accelerates mammary tumorigenesis in Wnt-1 transgenic mice and promotes chromosomal instability. Genes Dev. 9:882-95, 1995.
309. Taylor, I.C., Roy, S., Yaswen, P., Stampfer, M.R., Varmus, H.E. Mouse mammary tumors
express elevated levels of RNA encoding the murine homolog of SKY, a putative receptor tyrosine kinase. J Biol. Chem. 270:6872-80, 1995
310. Kaplan, K.B., Swedlow, J.R., Morgan, D.O., Varmus, H.E. c-Src enhances the spreading of
src -/- fibroblasts on fibronectin by a kinase-independent mechanism. Genes Dev. 9:1505-17, 1995.
311. Ranade, K., Hussussian, C.J., Sikorski, R.S., Varmus, H.E., Goldstein, A.M., Tucker, M.A.,
Serrano, M., Hannon, G.J., Beach, D., Dracopoli, N.C. Mutations associated with familial melanoma impair p16INK4 function. Nature Genetics 10:114-6, 1995.
312. Taylor, I.C., Roy, S., Varmus, H.E. Overexpression of the Sky receptor tyrosine kinase at the
cell surface or in the cytoplasm results in ligand-independent activation. Oncogene, 12:2619-26, 1995.
313. Varmus, H.E. Under the Influence: from provirus hypothesis to multistep carcinogenesis. In
The DNA Provirus: Howard Temin's Scientific Legacy, ASM Press, Wash, D.C., 1995. 314. Lowell, C.A., Niwa, M., Soriano, P., Varmus, H. Deficiency of the Hck and Src tyrosine
kinases results in extreme levels of extramedullary hematopoiesis. Blood 5:1780-92, 1996. 315. Leavitt, A.D., Robles, G., Alesandro, N., Varmus, H.E. Human immunodeficiency virus type
1 integrase mutants retain in vitro integrase activity yet fail to integrate viral DNA efficiently during infection. J. Virol. 70:721-8, 1996.
316. Godley, L.A., Kopp, J.B., Eckhaus, M., Paglino, J.J., Owens, J., Varmus, H.E. Wild-type p53
transgenic mice exhibit altered differentiation of the ureteric bud and possess small kidneys. Genes Dev. 10:836-50, 1996.
317. Broccoli, D., Godley, L.A., Donehower, L.A., Varmus, H.E., de Lange, T. Telemerase
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activation in mouse mammary tumors: lack of detectable telomere shortening and evidence for regulation of telomerase RNA with cell proliferation. Mol. Cell. Biol. 16:3765-72, 1996
318. Chen, X., Kang, H., Shen, L.X., Chamorro, M., Varmus, H. E., Tinoco, I. Jr. A characteristic
bent conformation of RNA pseudoknots promotes -1 frameshifting during translation of retroviral RNA. J. Mol. Biol. 260:479-83, 1996.
319. Donehower, L.A., Godley, L.A., Aldaz, C.M., Pyle, R., Shi, Y.P., Pinkel, D., Gray, J.,
Medina, D., Varmus, H.E. The role of p53 loss in genomic instability and tumor progression in a murine mammary cancer model. Prog Clin Biol Res. 395:1-11, 1996.
320. He, X., Saint-Jeannet, J.P., Wang, Y., Nathans, J., Dawid, I., Varmus, H. A member of the
frizzled protein family mediating axis induction by Wnt-5a. Science 275:1652-4, 1997. 321. Willert, K., Brink, M., Wodarz, A., Varmus, H., Nusse, R. Casein kinase 2 associates with
and phosphorylates dishevelled. EMBO J. 16:101-8, 1997. 322. Finch, P.W., He, X., Kelley, M.J., Uren, A., Schaudies, R.P., Popescu, N.C., Rudikoff, S.,
Aaronson, S.A., Varmus, H.E., Rubin, J.S. Purification and molecular cloning of a secreted, Frizzled-related antagonist of Wnt action. Proc. Natl. Acad. Sci. 94:6770-5, 1997
323. Jones, J.M., Attardi, L., Godley, L.A., Laucirica, R., Medina, D., Jacks, T., Varmus, H.E.,
Donehower, L.A. Absence of p53 in a mouse mammary tumor model promotes tumor cell proliferation without affecting apoptosis. Cell Growth Differ 8:829-38, 1997.
324. Saint-Jeannet, J.P., He, X., Varmus, H.E., Dawid, I.B. Regulation of dorsal fate in the
neuraxis by Wnt-1 and Wnt-3a. Proc. Natl. Acad. Sci. 94:13713-8, 1997 325. Schwartzberg, P.L., Xing, L., Hoffmann, O., Lowell, C.A., Garrett, L., Boyce, B.F., Varmus,
H. Rescue of osteoclast function by transgenic expression of kinase-deficient Src in src-/- mutant mice. Genes Dev. 11:2835-44, 1997.
326. Bates, P., Rong, L., Varmus, H.E., Young, J.A., and Crittenden, L.B. Genetic mapping of the
cloned subgroup A avain sarcoma and leukosis virus receptor gene to the TVA locus. J. Virol. 72:2505-8, 1998
327. Holland, E.C., Varmus, H.E. Basic fibroblast growth factor induces cell migration and
proliferation after glia-specific gene transfer in mice. Proc. Natl. Acad. Sci. 95:1218-23, 1998
328. Holland, E.C., Hively, W.P., DePinho, R.A., Varmus, H.E. A constitutively active epidermal
growth factor receptor cooperates with disruption of G1 cell-cycle arrest pathways to induce glioma-like lesions in mice. Genes Dev. 12:3675-85, 1998.
329. Holland, E.C., Hively, W.P., Gallo, V., Varmus, H.E. Modeling mutations in the G1 arrest
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pathway in human gliomas: overexpression of CDK4 but not loss on INK4a-ARF induces hyperploidy in cultured mouse astrocytes. Genes Dev. 12:3644-49, 1998.
330. Debnath, J., Chamorro, M., Czar, M.J., Schaeffer, E.M., Lenardo, M.J., Varmus, H.E.,
Schwartzberg, P.L. rlk/TXK encodes two forms of a novel cysteine string tyrosine kinase activated by src family kinases. Mol. Cell Biol. 19:1498-507, 1999.
331. Bocchinfuso, W.P., Hively, W.P., Couse, J.F., Varmus, H.E., Korach, K.S. A mouse
mammary tumor virus-wnt-1 transgene induces mammary gland hyperplasia and tumorigenesis in mice lacking estrogen receptor-α. Cancer Res. 59:1869-76, 1999.
332. Schaeffer, E.M., Debnath, J., Yap, G., McVicar, D., Liao, X.C., Littman, D.R., Sher, A.,
Varmus, H.E., Lenardo, M.J., Schwartzberg, P.L. Requirement for Tec kinases Rlk and Itk in T Cell receptor signaling and immunity. Science 284:638-41, 1999.
333. Zorn, A.M., Barish, G.D., Williams, B.O., Lavender, P., Klymkowsky, M.W., Varmus, H.E.
Regulation of Wnt signaling by Sox proteins: XSox17a/b and XSox3 interact physically with ß-catenin/TCF activity. Mol Cell 4:487-98, 1999.
334. Klymkowsky, M.W., Williams, B.O., Barish, G.D., Varmus, H.E., Vourgourakis, Y.E.
Membrane anchored plakoglobins have multiple mechanisms of action in Wnt signaling. Mol Biol Cell 10:3151-69, 1999.
335. Fisher, G.H., Orsulic, S., Holland, E., Hively, W.P., Li, Y., Lewis, B., Williams, B.O.,
Varmus, H.E. Development of a flexible and specific gene delivery system for production of murine tumor models. Oncogene 18:5253-60, 1999.
336. Li, Y., Hively, W.P., Varmus, H.E Use of MMTV-Wnt-1 transgenic mice for studying the
genetic basis of breast cancer. Oncogene 19:1002-9, 2000. 337. Williams, B.O., Barish, G.D., Klymkowsky, M.W., Varmus, H.E. A comparative evaluation
of ß-catenin and plakoglobin signaling activity. Oncogene 19(50)5720-8, 2000. 338. Xing, L., Venegas, A.M., Chen, A., Garrett-Beal, L., Boyce, B.F., Varmus, H.E.,
Schwartzberg, P.L. Genetic evidence for a role for Src family kinases in TNF family receptor signaling and cell survival. Genes Dev 15:241-53, 2001.
339. Li, Y., Podsypanina, K., Liu, X., Crane, A., Tan, L. K., Parsons, R., and Varmus, H. E.
Deficiency of Pten accelerates mammary oncogenesis in MMTV-Wnt-1 transgenic mice, BMC Mol Biol. 2:2, 2001.
340. Chamorro, M. Czar, M.J., Debnath, J., Cheng, G., Lenardo, M.J., Varmus, H.E.,
Schwartzberg, P.L. Requirements for activation and RAFT localization of the T-lymphocyte kinase Rlk/Txk BMC Immunology 2001 2:3, 2001.
341. Lewis, B.C., Chinnasamy, N., Morgan, R.A., and Varmus, H.E. Development of an Avian
Leukosis Sarcoma Virus Subgroup A Pseudotyped Lentiviral Vector. J of Virol 75:9339-44,
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2001. 342. Fisher, G.H, Wellen, S.H., Klimstra, D., Lenczowski, J.M., Tichelaar, J.W., Lizak, M.J,
Whitsett, J.A., Koretsky, A. and Varmus, H.E. Induction and apoptotic regression of lung adenocarcinomas by regulation of a K-Ras transgene in the presence and absence of tumor suppressor genes. Genes Dev 15:3249-62, 2001.
343. Orsulic, S., Li, Y., Soslow, R.A., Vitale-Cross, L.A., Gutkind, S.J., and Varmus, H.E.
Induction of ovarian cancer by defined multiple genetic changes in a mouse model system. Cancer Cell 1:53-62, 2002.
344. Schweizer, L. and Varmus, H.E. Wnt/Wingless signaling through ß-catenin requires function
of both LRP/Arrow and Frizzled classes of receptors. BMC Cell Biology 4:4, 2003. 345. Pao, W., Klimstra, D.S., Fisher, G.H., and Varmus, H.E. Use of avian retroviral vectors to
introduce transcriptional regulators into mammalian cells for analyses of tumor maintenance. Proc. Natl. Acad. Sci. 100:8764-9, 2003.
346. Cong, F., Zhang, J., Pao, W., Zhou, P. and Varmus, H.E. A protein knockdown strategy to
study the function of ß-catenin in tumorigenesis. BMC Molecular Biology, 4:10, 2003. 347. Li, Y., Welm, B., Podsypanina, K., Huang, S., Chamorro, M., Zhang, X., Rowlands, T.,
Egeblad, M., Cowin, P., Werb, Z., Tan, L.K., Rosen J.M., Varmus H.E. Evidence that transgenes encoding components of the Wnt signaling pathway preferentially induce mammary cancers from progenitor cells. Proc. Natl. Acad. Sci. 100:15853-8, 2003.
348. Lewis, B.C, Klimstra, D.S. and Varmus, H.E. The c-myc and PyMT oncogenes induce
different tumor types in a somatic mouse model for pancreatic cancer. Genes Dev. 17:3127-38, 2003.
349. Cong, F., Schweizer, L., Chamorro, M. and Varmus, H.E. Requirement for a nuclear function
of ß-catenin in Wnt signaling. Mol. Cell Biol. 23:8462-70, 2003. 350. Cong, F. and Varmus, H.E. Nuclear-cytoplasmic shuttling of Axin regulates subcellular
localization of ß-catenin. Proc. Natl. Acad. Sci. 101:2882-7, 2004. 351. Cong, F., Schweizer, L., and Varmus, H.E. Casein kinase Іε modulates the signaling
specificities of dishevelled. Mol. Cell. Biol. 24:2000-11, 2004. 352. Cong, F. Schweizer, L. and Varmus, H.E. Wnt signals across the plasma membrane to
activate the ß-catenin pathway by forming oligomers containing its receptors, Frizzled and LRP. Development 131:5103-15, 2004.
353. Chamorro, M., Schwartz, D.R., Vonica, A., Brivanlou, A.H., Cho, K.R., and. Varmus, H.E.
FGF-20 and DKK1 are transcriptional targets of beta-catenin and FGF-20 is implicated in cancer and development. EMBO J., 24:73 – 84, 2005.
354. Podsypanina, K., Li, Y., and Varmus, H. E. Evolution of somatic mutations in mammary
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tumors in transgenic mice is influenced by the inherited genotype. BMC Med 2:1 24, 2004. 355. Pao, W., Miller, V., Zakowski, M., Doherty, J., Politi, K., Sarkaria, I., Singh, B., Heelan, B.,
Rusch, V., Fulton, L., Mardis, E., Kupfer, D, Wilson, R., Kris, M., and Varmus, H.E. EGF receptor gene mutations are common in lung cancers from "never smokers" and are associated with sensitivity of tumors to gefitinib and erlotinib. Proc. Natl. Acad. Sci. 101:13306-11, 2004.
356. Lewis, B.C., Klimstra, D.S., Socci, N.D., Koutcher, J.A. and Varmus, H.E. The absence of
p53 promotes metastasis in a novel somatic mouse model for hepatocellular carcinoma. Mol Cell Biol 25:4 1228-37, 2005.
357. Pao, W., Wang, T.Y., Riely, G.J., Miller, V.A., Pan, Q., Ladanyi, M., Zakowski. M.F.,
Heelan, R.T., Kris, M.G., Varmus, H.E. KRAS mutations and primary resistance of lung adenocarcinomas for gefitinib or erlotinib. PLoS Medicine 2:1 e17, 2005.
358. Pao, W., Miller, V.A., Politi, K., Riely, G.J., Somwar, R., Zakowski. M.F., Kris, M.G.,
Varmus, H.E. Acquired resistance of lung adenocarcinomas to gefitinib or erlotinib is associated with a second mutation in the EGFR kinase domain. PLoS Med 2:3 e73, 2005.
359. Huang, S., Li, Y., Chen,Y., Podsypanina, K., Chamorro, M., Olshen, A.B., Desai, K.B., Tann,
A., Petersen, D., Green, J.E., and Varmus, H.E. Changes in gene expression during the development of mammary tumors in MMTV-Wnt-1 transgenic mice. BMC Mol Biol 6:10, R84, 2005.
360. Carter, T.A., Wodicka, L.M., Shah, N.P., Velasco, A.M., Fabian, M.A., Treiber, D.K.,
Milanov, Z.V., Atteridge, C.E., Biggs III, W.E., Edeen, P.T., Floyd, M., Ford J.M., Grotzfeld. R.M., Herrgard, S., Insko, D.E., Mehta, S.A., Patel, H.K., Pao, W., Sawyers , C.L., Varmus, H.E., Zarrinkar, P.P., and Lockhart, D.J. Inhibition of drug-resistant mutants of ABL, KIT, and EGF receptor kinases. Proc Natl Acad Sci U S A 102:11011–6, 2005.
361. Varmus, H., Pao, W., Politi, K., Podsypanina, K., Du, Y.-C.N. Oncogenes Come of Age.
CSHSQB 70:1-9, 2005. 362. Politi, K., Zakowski, M.F., Fan, P., Schonfeld, E.A. Pao, W. and Varmus, H.E. Lung
adenocarcinomas induced in mice by mutant EGF receptors found in human lung cancers respond to a tyrosine kinase inhibitor or to down-regulation of the receptors. Genes Dev. 20:1496-1510, 2006.
363. Du Z, Podsypanina K, Huang S, McGrath A, Toneff MJ, Bogoslovskaia E, Zhang X, Moraes
RC, Fluck M, Allred DC, Lewis MT, Varmus HE, Li Y. Introduction of oncogenes into mammary glands in vivo with an avian retroviral vector initiates and promotes carcinogenesis in mouse models. Proc Natl Acad Sci U S A. 46:17396-401, 2006.
364. Marks, J.L., McLellan, M.D., Zakowski, M.F., Lash, A.E., Kasai, Y., Broderick, S., Sarkaria,
I.S., Pham, D., Singh, B., Miner, T.L., Fewell, G.A., Fulton, L.L., Mardis, E.R., Wilson, R.K., Kris, M.G., Rusch, RW, Varmus, H.E., Pao, W. Mutational Analysis of EGFR and Related Signaling Pathway Genes in Lung Adenocarcinomas Identifies a Novel Somatic
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Kinase Domain Mutation in FGFR4. PLoS One 2: e426, 2007. 365. Du, Y-C.N., Lewis, B.C., Hanahan, D., and Varmus, H.E. Assessing Tumor Progression
Factors by Somatic Gene Transfer in a Mouse Model: Bcl-xL Promotes Islet Tumor Cell Invasion. PLoS Biology 5: e276, 2007
366. Weir, B.A., Woo, M.S., Getz, G., Perner, S., Ding, L., Beroukhim, R., Lin, W.M., Province,
M.A., Kraja, A., Johnson, L.A., Shah, K. Sato, M., Thomas, R.K., Barletta, J.A., Borecki, I.B., Broderick, S., Chang, A.C. Chiang, D.Y., Chirieac, L.R., Cho, J., Fujii, Y., Gazdar, A.F., Giordano, T., Greulich, H., Hanna, M., Johnson,B.E., Kris, M.G., Lash, A., Lin, L., Lindeman, N., Mardis, E.R., McPherson, J.D., Minna, J.D., Morgan, M.B., Nadel, M., Orringer, M.B., Osborne, J.R., Ozenberger, B., Ramos, A.H., Robinson, J., Roth, J.A., Rusch, V., Sasaki, H., Shepherd, F., Sougnez, C., Spitz, M.R., Tsao, M-S., Twomey, D., Verhaak, R.G.W., Weinstock, G.M., Wheeler, D.A., Winckler, W., Yoshizawa, A., Yu, S., Zakowski, M.F., Zhang, Q., Beer, D.G., Wistuba, I.I, Watson, M.A., Garraway, L.A., Ladanyi, M., Travis, W.D., Pao, W., Rubin, M.A., Gabriel, S.B., Gibbs, R.A., Varmus, H.E., Wilson, R.K., Lander E.S., & Meyerson, M. Characterizing the cancer genome in lung adenocarcinoma. Nature 450: 893-898 (2007)
367. K. Podsypanina, K.A. Politi, L.J. Beverly and H. E. Varmus. Oncogene cooperation in tumor
maintenance and tumor recurrence in mouse mammary tumors induced by Myc and mutant Kras. PNAS 105: 5242-5247 (2008)
368. Guha, U., Chaerkedy, R., Marimuthu, A., Patterson, A.S., Kashyap, M., Gowda, H., Sato, M.,
Bader, J., Lash, A.E., Minna, J.D., Pandey, A., Varmus, H.E.. Comparisons of tyrosine-phosphorylated proteins in immortalized human bronchial epithelial cells and lung adenocarcinoma cell lines expressing lung cancer-specific alleles of EGFR and KRAS. PNAS 105:14112-7 (2008)
369. Guha U, Chaerkady R, Marimuthu A, Patterson AS, Kashyap MK, Harsha HC, Sato M, Bader
JS, Lash AE, Minna JD, Pandey A, Varmus HE. Comparisons of tyrosine-phosphorylated proteins in immortalized human bronchial epithelial cells and lung adenocarcinoma cell lines expressing lung cancer-specific alleles of EGFR and KRAS. PNAS 105:14112-7 (2008)
370. K. Podsypanina, Y-C. N. Du, M. Jechlinger, L.J. Beverly, D. Hambardzumyan, and H.
Varmus. Seeding and Propagation of Untransformed Mouse Mammary Cells in the Lung. Science 321: 1841-1844 (2008)
371. Ding L, Getz G, Wheeler DA, Mardis ER, McLellan MD, Cibulskis K, Sougnez C, Greulich H, Muzny DM, Morgan MB, Fulton L, Fulton RS, Zhang Q, Wendl MC, Lawrence MS, Larson DE, Chen K, Dooling DJ, Sabo A, Hawes AC, Shen H, Jhangiani SN, Lewis LR, Hall O, Zhu Y, Mathew T, Ren Y, Yao J, Scherer SE, Clerc K, Metcalf GA, Ng B, Milosavljevic A, Gonzalez-Garay ML, Osborne JR, Meyer R, Shi X, Tang Y, Koboldt DC, Lin L, Abbott R, Miner TL, Pohl C, Fewell G, Haipek C, Schmidt H, Dunford-Shore BH, Kraja A, Crosby SD, Sawyer CS, Vickery T, Sander S, Robinson J, Winckler W, Baldwin J, Chirieac LR, Dutt A, Fennell T, Hanna M, Johnson BE, Onofrio RC, Thomas RK, Tonon G, Weir BA, Zhao X, Ziaugra L, Zody MC, Giordano T, Orringer MB, Roth JA, Spitz MR, Wistuba II, Ozenberger B, Good PJ, Chang AC, Beer DG, Watson MA, Ladanyi M, Broderick S,
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Yoshizawa A, Travis WD, Pao W, Province MA, Weinstock GM, Varmus HE, Gabriel SB, Lander ES, Gibbs RA, Meyerson M, Wilson RK. Somatic mutations affect key pathways in lung adenocarcinoma. Nature. 455(7216):1069-75 (2008)
372. Beverly, L. and Varmus, H. MYC-induced myeloid leukemogenesis is accelerated by all six
members of the anti-apoptotic BCL family. Oncogene 28, 1274 - 1279 (2009). NIHMSID: 99926
373. Asimakopoulos, F. and Varmus, H. Cell-specific Transduction of Prdm1-expressing Lineages
Mediated by a Receptor for Avian Leukosis Virus Subgroup B. J. Virol May;83 (10):4835-43. (2009) PMCID: PMC2682090
374. Jechlinger, M., Podsypanina, K., and Varmus, H. Regulation of transgenes in three-
dimensional cultures of primary mouse mammary cells demonstrates oncogene dependence and identifies cells that survive deinduction.. Genes Dev. Jul 15;23(14):1677-88. (2009) PMCID: PMC2714708
375. Du, Y.-C. N., Klimstra, D., and Varmus, H. Activation of PyMT in b cells induces
irreversible benign b-cell hyperplasia but oncogene-dependent acinar cell carcinomas when activated in pancreatic progenitor cells. PLoS ONE, 4, e6932 (2009) PMCID: PMC2758666
376. Somwar, R., Shum, D., Djaballah, H. and Varmus, H. Identification and preliminary characterization of novel small molecules that inhibit growth of human lung adenocarcinoma cells. J Biomol Screen. Dec;14(10):1176-84. Epub. (2009) PMCID: PMC2722688
377. Politi K, Fan PD, Shen R, Zakowski M, Varmus H. Erlotinib resistance in mouse models of epidermal growth factor receptor-induced lung adenocarcinoma. Dis Model Mech. 2010 Jan-Feb;3(1-2):111-9. Epub 2009 Dec 9. PMCID: PMC2806903
378. Barretina J, Taylor BS, Banerji S, Ramos AH, Lagos-Quintana M, Decarolis PL, Shah K,
Socci ND, Weir BA, Ho A, Chiang DY, Reva B, Mermel CH, Getz G, Antipin Y, Beroukhim R, Major JE, Hatton C, Nicoletti R, Hanna M, Sharpe T, Fennell TJ, Cibulskis K, Onofrio RC, Saito T, Shukla N, Lau C, Nelander S, Silver SJ, Sougnez C, Viale A, Winckler W, Maki RG, Garraway LA, Lash A, Greulich H, Root DE, Sellers WR, Schwartz GK, Antonescu CR, Lander ES, Varmus HE, Ladanyi M, Sander C, Meyerson M, Singer S. Subtype-specific genomic alterations define new targets for soft-tissue sarcoma therapy. Nat Genet. 2010 Aug. 42(8): 715-21. Epub 2010 July 4.
379. Somwar R, Erdjument-Bromage H, Larsson E, Shum D, Lockwood WW, Yang G, Sander C,
Ouerfelli O, Tempst PJ, Djaballah H,Varmus H. Superoxide dismutase 1 (SOD1) is a target for a small molecule identified in a screen for inhibitors of the growth of lung adenocarcinoma cell lines. PNAS 108: 16375-80 (2011)
380. Taguchi A, Politi K, Pitteri SJ, Lockwood WW, Faca VM, Kelly-Spratt K, Wong CH, Zhang
Q, Chin A, Park KS, Goodman G, Gazdar AF, Sage J, Dinulescu DM, Kucherlapati R, DePinho RA, Kemp CJ, Varmus HE, Hanash SM. Lung cancer signatures in plasma based on proteome profiling of mouse tumor models. Cancer Cell 20 (3): 289-299 (2011)
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381. Du YN, Chou CK, Klimstra DS, Varmus H. Receptor for hyaluronan-mediated motility isoform B promotes liver metastasis in a mouse model of multistep tumorigenesis and a tail vein assay for metastasis. PNAS 108: 16753-8 (2011)
382. Beverly LJ, Lockwood WW, Shah PP, Erdjument-Bromage H, Varmus H. Ubiquitination,
localization and stability of an anti-apoptotic BCL2-like protein, BCL2L10/BCLb, are regulated by Ubiquilin1. PNAS: 109: 119-26 (2012)
383. Nusse R and Varmus H. Three decades of Wnts: A personal perspective on how a scientific
field developed. The EMBO Journal 31: 2670-84 (2012) 384. Lockwood WW, Zejnullahu K, Bradner JE, Varmus H. Sensitivity of human lung
adenocarcinoma cell lines to targeted inhibition of BET epigenetic signaling proteins. PNAS 109: 19408-13 (2012)
385. Berger AH, Chen M, Morotti A, Janas JA, Niki M, Bronson RT, Taylor BS, Ladanyi M, Aelst
LV, Politi K, Varmus HE, Pandolfi PP. DOK2 inhibits EGFR-mutant lung adenocarcinoma. PLOS ONE 8: e79526 (2013)
386. de Bruin EC, Cowell C, Warne PH, Jiang M, Saunders RE, Melnick ME, Gettinger S,
Walther Z, Wurtz A, Heynen GJ, Heideman DAM, Gomez-Roman J, Garcia-Castano A, Gong Y, Ladanyi M, Varmus H, Bernards R, Smit EF, Politi K, Downward J. Reduced NF1 expression confers resistance to EGFR inhibition in lung cancer. Cancer Discovery 4(5): 605-19 (2014)
387. Song X, Fan P-D, Bantikassegn A, Guha U, Threadgill DW, Varmus H, Politi K. ERBB3-
independent activation of the PI3K pathway in EGFR-mutant lung adenocarcinomas. Cancer Research. 2015 Mar 15;75(6):1035-45.
388. Maity TK, Venugopalan A, Linnoila I, Cultraro CM, Giannakou A, Nemati R, Zhang X,
Webster JD, Ritt D, Ghosal S, Hoschuetzky H, Simpson RM, Biswas R, Politi K, Morrison DK, Varmus HE, Guha U. Loss of Mig6 accelerates initiation and progression of mutant epidermal growth factor receptor-driven lung adenocarcinoma. Cancer Discovery. 2015 May;5(5):534-49.
389. Unni AM, Lockwood WW, Zejnullahu K, Lee-Lin SQ, Varmus H. Synthetic lethality
explains mutually exclusive mutations of KRAS and EGFR in lung adenocarcinoma. Elife. 2015 Jun 5;4.
390. Varmus H, Kuehn BM. Varmus Returns to His Roots in Cancer Genetics. JAMA. 2015 Jul
7;314(1):13-4. Other publications by members of the Varmus Laboratory:
1. Carlson BA, Kwon SY, Chamorro M, Oroszlan S, Hatfield DL, Lee BJ.Transfer RNA modification status influences retroviral ribosomal frameshifting. Virology Mar 1;255(1):2-8, 1999.
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2. Orsulic S. An RCAS-TVA-based approach to designer mouse models. Mammalian
Genome 13;543-547, 2002.
3. Miller VA, Kris MG, Shah N, Patel J, Azzoli C, Gomez J, Krug LM, Pao W, Rizvi N, Pizzo B, Tyson L, Venkatraman E, Ben-Porat L, Memoli N, Zakowski M, Rusch V, Heelan RT. Bronchioloalveolar pathologic subtype and smoking history predict sensitivity to gefitinib in advanced non-small cell lung cancer (NSCLC). J Clin Oncol 22:1103-1109, 2004.
4. Ren G, Wang TY, Pan Q, Pao W, Huai J. Epidermal growth factor receptor mutations
detected in tumors from Chinese “never smokers” with lung adenocarcinoma. Chinese Med J; 118:769-771, 2005.
5. Zhang X, Podsypanina K, Huang S, Mohsin SK, Chamness GC, Hatsell S, Cowin P, Schiff
R, Li Y. Estrogen receptor positivity in mammary tumors of Wnt-1 transgenic mice is influenced by collaborating oncogenic mutations. Oncogene 24(26):4220-31, 2005.
6. Huang S, Podsypanina K, Chen Y, Cai W, Tsimelzon A, Hilsenbeck S, Li Y. Wnt-1 is
dominant over neu in specifying mammary tumor expression profiles. Technol Cancer Res Treat. Dec;5(6):565-71. 2006
7. Christiani DC, Pao W, DeMartini JC, Linnoila RI, Malkinson AM, Onn A, Politi KA, Sharp
M, Wong KK. BAC consensus conference, November 4-6, 2004: epidemiology, pathogenesis, and preclinical models. J Thorac Oncol. 1(9 Suppl):S2-7. (Review). 2006.
8. Riely, G.J., Politi, K.A., Miller, V A., Pao, W., Update on epidermal growth factor receptor
mutations in non-small cell lung cancer. Clin Cancer Res. 12(24):7232-41 (Review). 2006.
9. Gong Y, Somwar R, Politi K, Balak M, Chmielecki J, Jiang X, Pao W., Induction of BIM Is Essential for Apoptosis Triggered by EGFR Kinase Inhibitors in Mutant EGFR-Dependent Lung Adenocarcinomas. PLoS Med. 4(10):e294. 2007.
10. Regales L, Balak MN, Gong Y, Politi K, Sawai A, Le C, Koutcher JA, Solit DB, Rosen N,
Zakowski MF, Pao W., Development of new mouse lung tumor models expressing EGFR T790M mutants associated with clinical resistance to kinase inhibitors. PLoS ONE 2(8):e810. 2007.
11. Beverly LJ. Regulation of anti-apoptotic BCL2-proteins by non-canonical
interactions: the next step forward or twosteps back? J Cell Biochem. 2012 Jan;113(1):3-12. doi: 10.1002/jcb.23335. PMID:21898539
12. Lin L, Bass AJ, Lockwood WN, Wang Z, Silvers AL, Thomas DG, Chang AC, Lin J,
Orringer MB, Li W, Glover TW, Giordano TJ, Lam WL, Meyerson M, Beer DG. Activation of GATA binding protein 6 (GATA6) sustains oncogenic lineage-survival in esophageal adenocarcinoma. Proc Natl Acad Sci U SA. 2012 Mar 13;109(11):4251-6. doi: 10.1073/pnas.1011989109. Epub 2012 Feb 28. PMID:22375031
13. Lockwood WN, Wilson IM, Coe BP, Chari R, Pikor LA, Thu KL, Solis LM, Nunez Ml,
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Behrens C, Yee J, English J, Murray N, Tsao MS, Minna JD, Gazdar AF, Wistuba II, MacAulay CE, LamS, Lam WL. Divergent genomic and epigenomic landscapes of lung cancer subtypes underscore the selection of different oncogenic pathways during tumor development. PLoS One. 2012;7(5):e37775. doi: 10.1371/journal.pone.0037775. Epub 2012 May 21.PMID:22629454
14. Lockwood WN, Chandel SK, Stewart GL, Erdjument-Bromage H, Beverly LJ. The novel ubiguitin ligase complex. SCFCFbxw4), interacts with the COP9 signalosome in an F-box dependent manner, is mutated. lost and under-expressed in human cancers. PLoS One. 2013 May 2;8(5):e63610. doi: 10.1371/journal.pone.0063610.
15. Casson L, Howell L, Mathews LA, Ferrer M, Southall N, Guha R, Keller JM, Thomas C,
Siskind LJ, Beverly LJ. Inhibition of ceramide metabolism sensitizes human leukemia cells to inhibition of BCL2-Iike proteins. PLoS One. 2013;8(1):e54525. doi: 10.1371/journal.pone.0054525. Epub 2013 Jan 16. PMID:23342165
16. Furlan SN, Mandraju R, Brewer T, Roybal K, Troutman TO, Hu W, Palm NW, Unni A,
Pasare C. Enhancement of anti-tumor CD8 immunity by lgG1-mediated targeting of Fe receptors. MAbs. 2014 Jan 1;6(1):108-18. doi: 10.4161/mabs.27052. Epub 2013 Nov 11. PMID:24284965