Principal Investigator/Program Director (Last, First, Middle): Helfand, Stephen L. POSITION TITLE

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NAME Stephen L. Helfand POSITION TITLE Professor era COMMONS USER NAME Shelfand EDUCATION/TRAINING (Begin with baccalaureate or other initial professional education, such as nursing, and include postdoctoral
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NAME Stephen L. Helfand POSITION TITLE Professor era COMMONS USER NAME Shelfand EDUCATION/TRAINING (Begin with baccalaureate or other initial professional education, such as nursing, and include postdoctoral training.) DEGREE INSTITUTION AND LOCATION (if applicable) YEAR (s) FIELD OF STUDY SUNY Binghamton, Binghamton NY Stanford University, Stanford, CA B.Sc Biology Albert Einstein College of Medicine, Bronx, NY M.D Medicine Professional Experience 1980 Medical Intern, Internal Medicine, Montefiore Hospital and Medical Center 1981 Visiting Scholar, Neuroimmunology, Queens University Clinical Resident, Neurology, Massachusetts General Hospital, Clinical Fellow, Neurology, Harvard Medical School Postdoctoral Fellow, Dept. of Biological Sciences, Stanford University Research Associate, Dept. of Biological Sciences, Stanford University Associate Research Scientist, Dept. of Biology, Yale University Assistant Professor, Dept. of BioStructure and Function, Univ. Conn. Health Center 1997 Associate Professor, Dept. of BioStructure and Function, Univ. Conn. Health Center Director, Developmental Biology/Genetics Graduate Program, Univ. Conn. Health Center Associate Professor, Dept. of Genetics and Developmental Biology,Univ. Conn. Health Center Professor, Department of Genetics and Developmental Biology, Univ. Conn. Health Center Professor, Department of Molecular Biology, Cell Biology, and Biochemistry, Brown Univ. Fellowships, Awards, and Honors Woods Scholarship, Stanford University 1974 Henry Rushton Fairclough Prize for Classics at Stanford University 1975 Distinction, Departmental Honors (Biology) Stanford University 1975 NSF Undergraduate Fellowship (Advisor: N.K. Wessells) Stanford University 1976 Predoctoral Trainee Fellowship (Advisor: D.P. Purpura and N.K. Wessells) Stanford University and Albert Einstein College of Medicine 1977 Roche Award in Neuroscience; American Medical Student Association-National Research Meeting Award 1978 Albert Cass Scholarship for Research in Neuroimmunology 1979 Medical Research Council Fellowship in Neuroimmunology (Advisor: M.C. Raff), Department of Zoology, University College, London 1981 Queen's University Development Fund Award-Neuroimmunology (Advisor: J.C. Roder), Department of Biology, Queen's University, Kingston, Ontario 1983 Boston Society of Psychiatry and Neurology Sandoz Award for Medical Research, Department of Neurology, Massachusetts General Hospital NIH Postdoctoral Fellowship, Stanford University NIH Physician Scientist Award, Stanford University and Yale University 1986 Diplomate, American Board of Psychiatry and Neurology 1990 University of Connecticut Research Initiation Program Grant 1991 American Cancer Society Institutional Research Grant Award 1991 University of Connecticut Faculty Research Grant 1992 American Federation of Aging Research Award PHS 398/2590 (Rev. 09/04) Page 6 1993 Sandoz Award for Gerontological Research 1993 University of Connecticut Faculty Research Grant The Patrick & Catherine Weldon Donaghue Medical Research Foundation Award, mentor for Dr. Blanka Rogina Gordon Conference Travel Award Summer Training Course in Experimental Aging Research, National Institutes on Aging 1995 American Federation of Aging Research Fellowship, mentor for Mr. Timothy Kuwada American Federation of Aging Research Award 1995 Nathan W. and Margaret T. Shock Aging Research Foundation Award 1998 University of Connecticut Faculty Research Grant The Donaghue Foundation Investigator Program for Health-Related Research Award 2000 Burroughs Wellcome Travel Grant to study in UK Ellison Medical Foundation Senior Investigator Award 2003 American Federation of Aging Research Fellowship, mentor for Mr. Brian Silvia American Federation of Aging Research Fellowship, mentor for Mr. Sanchez-Blanco R-37 NIH MERIT Award, National Institutes on Aging KO1 mentor for Dr. Yih-Woei Fridell American Federation of Aging Research Fellowship, mentor for Postdoctoral Fellowship for Dr. Johannes Bauer NIA K99 mentor for Dr. Johannes Bauer 2007 Keynote speaker at Biology of Aging Gordon Conference, Switzerland Glenn Award for Research in Biological Mechanisms of Aging Glenn Award for Research in Biological Mechanisms of Aging Publications Helfand, S.L., Smith, G.A., and Wessells, N.K. Survival and development in culture of dissociated parasympathetic neurons from ciliary ganglia. Dev. Biol. 1976; 50: Helfand, S.L., Riopelle, R.J., and Wessells, N.K. Non-equivalence of conditioned medium and nerve growth factor for sympathetic, parasympathetic and sensory neurons. Exp. Cell Res. 1978; 113: Helfand, S.L., Werkmeister, J., and Roder, J.C. The relationship between target cell binding, chemiluminescence and cytolysis. J. Exp. Med. 1982; 156: Roder, J.C., Helfand, S.L., Werkmeister, J., McGarry, R., Beaumont, T.J., and Duwe, A. Oxygen intermediates are triggered early in the cytoytic pathway of human NK cells. Nature 1982; 298: Werkmeister, J.A., Helfand, S.L., Rubin, P., Haliotis, T., Pross, H., and Roder, J.C. Tumor cell differentiation modulates susceptibility to natural killer cells. Cell. Immun. 1982; 69: Helfand, S.L., Werkmeister, J., and Roder, J.C. The role of free oxygen radicals in the activation of the NK cytolytic pathway. In: Herberman, R., editor. NK Cells and Other Natural Effector Cells. New York: Academic Press; 1982 p Werkmeister, J., Helfand, S.L., Haliotis, T., Pross, H., and Roder, J.C. (1982) Specificity of natural killer (NK) cells: nature of target cell structure. In: Herberman, R., editor, NK Cells and Other Effector Cells. New York: Academic Press; 1982 p Roder, J.C., Todd, R.F., Rubin, P., Haliotis, T., Helfand, S.L., Werkmeister, J., Pross, H.F., Boxer, L.A., Schlossman, S.F., and Fauci, A.S. The Chediak-Higashi gene in humans III. Studies on the mechanism of NK impairment. Clinical Exp. Immuno. 1983; 51: Helfand, S.L., Werkmeister, J., Pross, H., and Roder, J.C. Oxygen intermediates are required for interferon activation of NK cells. J. Interferon Res. 1983; 2: PHS 398/2590 (Rev. 09/04) Page 7 Werkmeister, J., Helfand, S.L., Roder, J., and Pross, H. The chemiluminescence response of human natural killer cels. II. Association of a decreased response with low natural killer activity. Eur. J. Immunol. 1983; 13: McGarry, R.C., Helfand, S.L., Quarles, R.H., and Roder, J.C. Recognition of myelin-associated gylcoprotein by the monoclonal antibody HNK-1. Nature 1983; 306: Goodman, C.S., Bastiani, M.M., Doe, C.Q., du Lac, S., Helfand, S.L., Kuwada, K.Y., Thomas, J.B. Neuronal recognition during development: cellular and molecular approaches. Science 1984; 225: Bastiani, M.J., Doe, C.Q., Helfand, S.L., and Goodman, C.S. Neuronal specificity and growth cone guidance in grasshopper and Drosophila embryos. Trends in Neuroscience 1985; 84: Beachy, P.A., Helfand, S.L. Hogness, D.S. Segmental distribution of bithorax complex proteins during Drosophila development. Nature 1985; 313: Hogness, D.S., Lipshitz, H.D., Beachy, P.A., Peattie, D.A., Saint, R.A., Goldschmidt-Clermont, M., Harte, P.J., Gavis, E.R., and Helfand, S.L. Regulation and products of the Ubx domain of the bithorax complex. Cold Spring Harbor Symp. Quant. Biol. 1985; 50: Helfand, S.L. and Carlson, J. Isolation and characterization of an olfactory mutant in Drosophila with a chemically specific defect. Proc. Natl. Acad. Sci. USA 1989; 86: Woodward, C., Huang,T., Sun, H., Helfand, S.L., and Carlson, J. Genetic analysis of olfactory behavior in Drosophila: A new screen yields the ota mutants. Genetics. 1989; 123: McKenna, M., Monte, P., Helfand, S.L., Woodard, C., and Carlson, J. A novel chemosensory response in Drosophila and the isolation of the acj mutations which affect it. Proc. Natl. Acad. Sci. USA 1989; 86: Irvine, K.D., Helfand, S.L. and Hogness, D.S. The large upstream control region of the Drosophila homeotic gene Ultrabithorax. Development 1991; 111: *Helfand, S.L., Blake,K.J., Rogina, B., Stracks, M.D., Centurion, A. and Naprta, B. Temporal patterns of gene expression in the antenna of the adult Drosophila melanogaster. Genetics 1995; 140: Blake, K.J., Rogina, B., Centurion, A. and *Helfand, S. L. Changes in gene expression during post-eclosional development in the olfactory system of Drosophila melanogaster. Mechanisms of Development 1995; 52: Rogina, B. and *Helfand, S. L. Regulation of gene expression is linked to life span in the adult Drosophila. Genetics 1995; 141: Blake, K. J., Hoopengardner, B., Centurion, A. and *Helfand, S. L. A molecular marker shows that adult maturation is independent of the rate of pre-adult development in Drosophila melanogaster. Developmental Genetics 1996; 18: *Helfand, S. L. and Naprta, B.. The expression of a reporter protein, ß-galactosidase, is preserved during maturation and aging in some cells of the adult Drosophila melanogaster. Mechanisms of Development 1996; 55: Rogina, B. and *Helfand, S. L. Timing of expression of a gene in the adult Drosophila is regulated by mechanisms independent of temperature and metabolic rate. Genetics 1996; 143: PHS 398/2590 (Rev. 09/04) Page 8 Rogina, B. and *Helfand, S. L. Spatial and temporal pattern of expression of the wingless and engrailed genes in the adult antenna is regulated by age-dependent mechanisms. Mechanisms of Development 1997; 63: Rogina, B., Benzer, S., and *Helfand, S. L. Drosophila drop-dead mutations accelerate the time course of agerelated markers. Proc. Natl. Acad. Sci. (USA) 1997; 94: Rogina, B., Vaupel, J. W., Partridge, L., *Helfand, S. L. Regulation of gene expression is preserved in aging Drosophila melanogaster. Current Biology 1998; 8: Rogina, B. and *Helfand, S.L. Cu, Zn superoxide dismutase deficiency accelerates the time course of an agerelated marker in Drosophila melanogaster. Biogerontology 2000; 1: Rogina, B. Reenan, R. A., Nilsen S. P. and *Helfand, S. L. Extended life-span conferred by cotransporter gene mutations in Drosophila. Science 2000; 290: *Helfand, S. L. and Rogina, B. Regulation of gene expression during aging. In: Hekimi, Siegfried, editor. Molecular Genetics of Aging: Results and Problems in Cell Differentiation, vol. 29. Germany: Springer-Verlag; p Hoopengardner, B and *Helfand, S. L. Temperature Compensation and Temporal Expression Mediated by an Enhancer Element in Drosophila. Mechanisms of Development 2002; 110: Knauf, F., Rogina, B., Jiang, Z., Aronson, P. A., and *Helfand, S. L. Functional Characterization and Immunolocalization of the Novel Transporter Encoded by the Life-Extending Gene Indy. Proc. Natl. Acad. Sci. (USA) 2002; 99: Rogina, B., Helfand, S. L. and Frankel, S. Longevity regulation by Drosophila Rpd3 deacetylase and caloric restriction. Science 2002; 298: *Helfand, S. L. Chaperones Take Flight. Science, 2002, 295: *Helfand, S. L. and Rogina, B. Molecular genetics of aging: Is this the end of the beginning? BioEssays 2003; 25: *Helfand, S. L. and Rogina, B. Genetics of aging in the fruit fly, Drosophila melanogaster. Annual Review of Genetics 2003; 37: *Helfand, S. L. and Inouye, S. L. Aging, life span, genetics and the fruit fly. Clinical Neuroscience Research 2003; 2: Butler, R. N., Austad, S. N., Barzilai, N., Braun, A., Helfand, S.L., Larsen, P. L., McCormick, A. M., Perls, T. T., Shuldiner, A. R., Sprott, R. L., and Warner, H. R. Longevity genes: from primitive organisms to humans. J Gerontol A Biol Sci Med Sci. 2003; 7: *Helfand, S. L. and Rogina, B. From genes to aging in the Drosophila. In: Hall, J.C., Dunlap, J. C., Friedman, T., editors. Advances in Genetics. San Diego: Academic Press; 2003 vol. 49, p *Helfand, S. L. and Rogina, B. Genetics of aging in the fruit fly, Drosophila melanogaster. Annual Review of Genetics 2003; 37: Marden, J.H., Rogina, B., Montooth,K.L. and *Helfand, S. L. Conditional tradeoffs between aging and organismal performance of Indy long-lived mutant flies.. Proc. Natl. Acad. Sci. (USA) 2003; 100: PHS 398/2590 (Rev. 09/04) Page 9 Fridell, Y-W., Sánchez-Blanco, Silvia, B. A. and *Helfand, S. L. Functional Characterization of a Drosophila Mitochondrial Uncoupling Protein. Journal of Bioenergetics and Biomembranes 2004; 36 (3): Woods, J, Rogina, B, Lavu, S., Howitz, K., *Helfand, S. L, *Tatar, M., and *Sinclair, D. (2004) Sirtuin activators mimic calorie restriction and delay aging in metazoans. Nature 2004; 430 (7000): Bauer, J., Goupil, S., Garber, G., and *Helfand, S. L. An accelerated assay for the identification of life span extending interventions in Drosophila melanogaster. Proc. Natl. Acad. Sci. (USA) 2004; 101: Rogina, B. and *Helfand, S. L. Sir2 mediates longevity in the fly through a pathway related to calorie restriction. Proc. Natl. Acad. Sci. (USA) 2004; 101: Fridell, Y-W, Sanchez-Blanco, A., Silvia, B. and *Helfand, S. L. Targeted Expression of the Human Uncoupling Protein 2 (hucp2) to Adult Neurons Extends Life Span in the Fly. Cell Metabolism 2005; 1: Zheng, J-Y, Mutcherson, R, and *Helfand, S. L. Calorie restriction delays lipid oxidative damage in Drosophila melanogaster. Aging Cell 2005; 4: Bross, TG, Rogina, B, and *Helfand SL. (2005) Behavioral, physical, and demographic changes in Drosophila populations through dietary restriction. Aging Cell. 4: (Cover picture) Bauer JH, Poon PC, Glatt-Deeley H, Abrams JM, and *Helfand S.L. (2005) Neuronal Expression of p53 Dominant-Negative Proteins in Adult Drosophila melanogaster Extends Life Span. Curr Biol. 15: Sanchez-Blanco A, Fridell YW, and *Helfand SL (2006) Involvement of Drosophila Uncoupling Protein 5 in Metabolism and Aging. Genetics 172:1-12. Knauf F, Mohebbi N, Teichert C, Herold D, Rogina B, Helfand SL, Gollasch M, Luft FC, and Aronson, PA. (2006) The life-extending gene Indy encodes an exchanger for Krebs-cycle intermediates. Biochemical Journal 397: Bauer, JH and *Helfand, S. L. The humble fly: what a model system can reveal about the human biology of aging. (2006) Rhode Island Medical Journal, 89(9): Bauer, JH and *Helfand, SL. (2006) New tricks of an old molecule: lifespan regulation by p53. Aging Cell 5: Bauer JH, Chang C, Morris SNS, Hozier S, Andersen S, Waitzman JS, *Helfand S.L. (2007) Expression of dominant-negative Dmp53 in the adult fly brain inhibits insulin signaling. Proc. Natl. Acad. Sci. (USA), 14; 104(33): Aug 8; [Epub ahead of print] *Helfand, S. L., Bauer, J. H. and Wood, J. G. (2008) Calorie Restriction in lower organisms in Molecular Biology of Aging edited by L. Guarente, L. Partridge, and D. Wallace. Cold Spring Harbor Laboratory Press, New York. Bauer J.H., Morris SNS, Chang C, Flatt T, Wood JG and *Helfand SL. (2009) dsir2 and Dmp53 interact to mediate aspects of CR-dependent life span extension in D. melanogaster. Aging 1: Neretti N, Wang PY, Brodsky AS, Nyguyen HH, White KP, Rogina B, *Helfand SL. (2009) Long-lived Indy induces reduced mitochondrial reactive oxygen species production and oxidative damage. Proc Natl Acad Sci U S A Jan 21. [Epub ahead of print] Wang PY, Neretti N, Whitaker R, Hosier S, Chang C, Lu D, Rogina B, *Helfand SL. (2009) Long-lived Indy and calorie restriction interact to extend life span. Proc Natl Acad Sci U S A Jun 9;106(23): Epub 2009 May 22. PHS 398/2590 (Rev. 09/04) Page 10 Fridell, Y-W, Hoh, M, Kreneisz, Orsolya, Hosier, S, Chengyi, C, Scantling D, Mulkey, D and *Helfand SL. Increased Uncoupling Protein (UCP) activity in Drosophila Insulin-Producing neurons attenuates Insulin signaling and extends lifespan. Aging (2009). Bauer, JH and *Helfand, SL. (2009) Sir2 and longevity: the p53 connection. Cell Cycle. Jun 15;8(12):1821. Epub 2009 Jun 15. * Contributing/Corresponding Author Research Projects Current 1R01 AG Helfand (PI) 9/30/05-6/30/10 Oxidative Damage, Aging and Life Span The major goals are to understand the role of oxidative damage in aging and life span. 2R37 AG16667 Helfand (PI) 3/31/09-3/31/14 Single Gene Mutants that Confer Longevity in Drosophila The major goals are to study the molecular genetic mechanisms of the Indy gene. 2R37 AG16667 Helfand (PI) ARRA Supplement 7/31/09-8/1/11 Single Gene Mutants that Confer Longevity in Drosophila The major goals are to study the molecular genetic mechanisms of the Indy gene. 2R37 AG16667 Helfand (PI) Underrepresented minority supplement 7/1/09-6/1/10 Single Gene Mutants that Confer Longevity in Drosophila The major goals are to study the molecular genetic mechanisms of the Indy gene. 1R01 AG24353 Helfand (PI) 8/1/09-7/31/14 Control of Gene Expression and Life Span The major goals are study the role of Rpd3 and Sir2 in life span determination. Glenn Award for Research in Biological Mechanisms of Aging Helfand (PI) 9/09-9/11 The goal of this project is to use biomarkers to identify new longevity mutants in Drosophila. Completed University of Connecticut Research Initiation Program Grant, PI, Molecular genetics of olfaction American Cancer Society Institutional Research Grant Award, PI, Genetic Studies of Signal Transduction in Olfaction. PHS 398/2590 (Rev. 09/04) Page 11 University of Connecticut Faculty Research Award, PI, Molecular and Genetic Analysis of Olfaction National Science Foundation, PI, Molecular and Genetic Analysis of Olfaction in Drosophila American Federation for Aging Research, PI, Regulation of Gene Expression During Aging Sandoz Foundation for Gerontological Research, PI, Gene regulation during aging University of Connecticut Faculty Research Award, PI, Regulation of Gene Expression During Aging American Federation for Aging Research, PI, Control of Gene Expression During Aging The Patrick & Catherine Weldon Donaghue Medical Research Foundation Award, Postdoctoral Mentor for Dr. Blanka Rogina 1995 Gerontology Society of America, Nathan W. and Margaret T. Shock Aging Research Foundation Award University of Connecticut Interim HCRAC Award, PI, Genetics of Aging in the Olfactory System of Drosophila Melanogaster RO3, PI Burroughs Wellcome Research Travel Grant, PI , PI, Single Gene Mutant that Confer Longevity in Drosophila The Patrick & Catherine Weldon Donaghue Medical Research Foundation, PI, Molecular Genetics of Aging Ellison Medical Foundation, PI, Isolation of Single Gene Mutations in Drosophila Melanogaster Glenn Award for Research in Biological Mechanisms of Aging, PI , Co-PI, Molecular Genetics of Caloric Restriction in Aging Flies , PI, MERIT Award, Single Gene Mutant that Confer Longevity AG , PI, Control of Gene Expression and Life Span AG24353 PHS 398/2590 (Rev. 09/04) Page 12
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