Carol W. Greider

Born in San Diego, California, Carol Greider discovered the first definitive indications of a new enzyme, originally called Tetrahymena telomere.  She was at University of California, Berkley as a first-year graduate student during that time.  She also made some significant discoveries at University of Göttingen.

She was awarded the 2009 Nobel Prize for Physiology or Medicine, along with Blackburn and Jack W. Szostak for their discovery that telomeres are protected from progressive shortening by the enzyme telomerase.

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Biologist BIOLOGIST (Molecular Biologist)
Carol Greider
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  1. Dickson Prize 2006
  2. Paul Ehrlich and Ludwig Darmstaedter Prize 2009 (with Elizabeth H. Blackburn)
  3. Horwitz Prize 2007 (with Elizabeth H. Blackburn and Joseph Gall)
  4. Lounsbery Award 2003
  5. Gairdner Foundation International Award 1998 (with Elizabeth H. Blackburn)
  6. Lasker Award 2006 (with Elizabeth H. Blackburn and Jack W. Szostak)
  7. Nobel Prize for Medicine 2009 (with Elizabeth H. Blackburn and Jack W. Szostak)
  8. Pew Scholar 1990
  9. Wiley Prize in Biomedical Sciences 2006 (with Elizabeth H. Blackburn)

1. Greider, C.W., and Blackburn, E.H. (1985). Identification of a specific telomere terminal transferase activity in Tetrahymena extracts. Cell 43: 405–413.
2. Greider, C.W. and Blackburn, E.H. (1987). The telomere terminal transferase of Tetrahymena is a ribonucleoprotein enzyme with two kinds of primer specificity. Cell 51: 887–898.
3. Greider, C.W., and Blackburn, E.H. (1989). A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis. Nature 337: 331–337.
4. Harley, C.B., Futcher, A.B., and Greider, C.W. (1990). Telomeres shorten during ageing of human fibroblasts. Nature 345: 458–460.
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5. Harrington, L.A., and Greider, C.W. (1991). Telomerase primer specificity and chromosome healing. Nature 353: 451–454.
6. Greider, C.W. (1991). Telomerase is processive. Mol. Cell Biol. 11: 4572–4580.
7. Allsopp, R.C., Vaziri, H., Patterson, C., Goldstein, S., Younglai, E.V., Futcher, A.B., Greider, C.W., and Harley, C.B. (1992). Telomere length predicts the replicative capacity of human fibroblasts. Proc. Natl. Acad. Sci. U.S.A. 89: 10114–10118.
8. Levy, M.Z., Allsopp, R.C., Futcher, A.B., Greider, C.W., and Harley, C.B. (1992). Telomere end replication problem and cell aging. J. Mol. Biol. 225: 951–960.
9. Counter, C.M., Avilion, A.A., LeFeuvre, C.E., Stewart, N.G., Greider, C.W., Harley, C.B., and Bacchetti, S. (1992). Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity. EMBO J. 11: 1921–1929.
10. Avilion, A.A., Harrington, L.A., and Greider, C.W. (1992). Tetrahymena telomerase RNA levels increase during macronuclear development. Dev. Genet. 13: 80–86.
11. Collins, K. and Greider C.W. (1993). Tetrahymena telomerase catalyzes nucleolytic cleavage and nonprocessive elongation. Genes Dev. 7: 1364-1376.
12. Prowse, K.R., Avilion, A.A., and Greider, C.W. (1993). Identification of a nonprocessive telomerase activity from mouse cells. Proc. Natl. Acad. Sci. U.S.A. 90: 1493-1497.
13. Mantell, L.L., and Greider, C.W. (1994). Telomerase activity in germline and embryonic cells of Xenopus. EMBO J. 13: 3211-3217.
14. Autexier, C., and Greider, C.W. (1994). Functional reconstitution of wild-type and mutant Tetrahymena telomerase. Genes Dev. 8: 563-575.
15. Collins, K., and Greider, C.W. (1995). Utilization of ribonucleotides and RNA primers by Tetrahymena telomerase. EMBO J. 14: 5422-5432.
16. Autexier, C., and Greider, C.W. (1995). Boundary elements of the Tetrahymena telomerase RNA template and alignment domains. Genes & Dev. 9: 2227-2239.
17. Blasco, M.A., Funk, W., Villeponteau, B., and Greider, C.W. (1995). Functional characterization and developmental regulation of the mouse telomerase RNA. Science 269: 1267-1270.
18. Feng, J., Funk, W.D., Wang, S.-S., Weinrich, S.L., Avilion, A.A., Chiu, C.-P., Adams, R.R., Chang, E., Allsopp, R.C., Yu, J., Le, S., West, M.D., Harley, C.B., Andrews, W.H., Greider, C.W., and Villeponteau, B. (1995). The RNA component of human telomerase. Science 269: 1236-1241.
19. Collins, K., Kobayashi, R., and Greider, C.W. (1995). Purification of Tetrahymena telomerase and cloning of genes encoding the two protein components of the enzyme. Cell 81: 677-686.
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20. Prowse, K.R., and Greider, C.W. (1995). Developmental and tissue specific regulation of mouse telomerase and telomere length. Proc. Natl. Acad. Sci. U.S.A. 92: 4818-4822.
21. Harrington, L.A., Hull, C., Crittenden, J., and Greider, C.W. (1995). Gel shift and UV cross-linking analysis of Tetrahymena telomerase. J. Biol. Chem. 270: 8893-8901.
22. Blasco, M., Rizen, M., Greider, C.W., and Hanahan, D. (1996). Differential regulation of telomerase activity and telomerase RNA during multi-stage tumorigenesis. Nat. Genet. 12: 200-204.
23. Avilion, A.A., Piatyszek, M. A., Gupta, J., Shay, J.W., Bacchetti, S., and Greider, C.W. (1996). Human telomerase RNA and telomerase activity in immortal cell lines and tumor tissues. Cancer Res. 56: 645-650.
24. Buchkovich, K.J., and Greider, C.W. (1996). Telomerase regulation during entry into the cell cycle in normal human T cells. Mol. Biol. Cell 7: 1443-1454.
25. Autexier, C., Pruzan, R., Funk, W., and Greider, C.W. (1996). Reconstitution of human telomerase activity and identification of a minimal functional region of the human telomerase RNA. EMBO J. 15: 5928-5935.
26. Blasco, M.A., Lee, H.-W., Hande, P.M., Samper, E., Lansdorp, P.M., DePinho, R.A., and Greider, C.W. (1997). Telomere shortening and tumor formation by mouse cells lacking telomerase RNA. Cell 91: 25-34.
27. Hinkley, C.S., Blasco, M.A., Funk, W.D., Feng, J., Villeponteau, B., Greider, C.W., and Herr, W. (1998). The mouse telomerase RNA 5’-end lies just upstream of the telomerase template sequence. Nucleic Acids Res. 26: 532-536.
28. Le, S., Zhu, J.J., Anthony, D.C., Greider, C.W., and Black, P.M. (1998). Telomerase activity in human gliomas. Neurosurgury 42: 1120-1125.
29. Autexier, C.A., and Greider, C.W. (1998). Mutational analysis of the Tetrahymena telomerase RNA: Identification of residues affecting telomerase activity in vitro. Nucleic Acids Res. 26: 787-795.
30. Lee, H.-W., Blasco, M.A., Gottlieb, G.J., Horner, J.W., Greider, C.W., and DePinho, R.A. (1998). Essential role of mouse telomerase in highly proliferative organs. Nature 392: 569-574.
31. Rudolph, K.L., Chang, S., Lee, H.-W., Blasco, M., Gottlieb, G., Greider, C.W., and DePinho, R.A. (1999). Longevity, stress response, and cancer in aging telomerase deficient mice. Cell 96: 701-712.
32. Le, S., Moore, J.K., Haber, J.E., and Greider, C.W. (1999). RAD51 and RAD50 define two pathways that collaborate to maintain telomeres in the absence of telomerase. Genetics 152: 143-152.
33. Greenberg, R., Chin, L., Femino, A., Lee, K.-H., Gottlieb, G., Singer, R., Greider, C.W., and DePinho, R.A. (1999). Short telomeres impair tumorigenesis in the INK4aD2/3-/- mTR-/- cancer-prone mouse. Cell 97: 515-525.
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34. Chin, L., Artandi, S., Shen, Q., Tam, S., Lee, S.-L., Gottlieb, G., Greider, C.W., and DePinho, R.A. (1999). p53 deficiency rescues the adverse effects of telomere loss in vivo and cooperates with telomere dysfunction to accelerate carcinogenesis. Cell 97: 527-538.
35. Hemann, M.T., and Greider, C.W. (1999). G-strand overhangs on telomeres in telomerase deficient mouse cells. Nucleic Acids Res. 27: 3964-3969.
36. Le, S., Sternglanz, R., and Greider, C.W. (2000). Identification of two RNA binding proteins associated with human telomerase RNA. Mol. Biol. Cell. 11: 999-1010.
37. Chen, J.-L., Blasco, M., and Greider, C.W. (2000). A Secondary structure of vertebrate telomerase RNA. Cell 100: 503-514.
38. Hemann, M. T., and Greider, C. W. (2000). Wild derived inbred mouse strains have short telomeres, Nucleic Acids Res. 28: 4474-4478.
39. Chen, Q., IJpma, A., and Greider, C. W. (2001). Two survivor pathways that allow growth in the absence of telomerase are generated by distinct telomere recombination events. Mol. Cell. Biol. 21: 1819-1827.
40. Hemann, M. T., Rudolph, L., Strong, M., DePinho, R. A., Chin, L., and Greider, C. W. (2001). Telomere dysfunction triggers developmentally regulated germ cell apoptosis. Mol. Biol. Cell. 12: 2023-2030.
41. Hackett, J., Feldser, D.M., and Greider, C. W. (2001). Telomere dysfunction increases mutation rate and genomic instability. Cell 106: 275-286.
42. Hemann, M. T., Strong, M., Hao, L.-Y., and Greider, C. W. (2001). The shortest telomere, not average telomere length, is critical for cell viability and chromosome stability. Cell 107: 66-77.
43. Mason, D., Autexier, C., and Greider, C. W. (2001). Tetrahymena proteins p80 and p95 are not core telomerase components. Proc. Natl. Acad. Sci. U.S.A. 98: 12368-12373.
44. Chen, J.-L., Opperman, K. K., and Greider, C. W. (2002). A critical stem loop structure in the CR4-CR5 domain of mammalian telomerase RNA. Nucleic Acids Res. 30: 592-597.
45. Hathcock, K. S., Hemann, M. T., Opperman, K. K., Strong, S. A., Greider, C. W., and Hodes, R. J. (2002). Haploinsufficiency of mTR results in defects in telomere elongation. Proc. Natl. Acad. Sci. U.S.A. 99: 3591-3596.
46. IJpma, A., and Greider, C.W. (2003). Short telomeres induce a DNA damage response in S. cerevisiae. Mol. Biol. Cell 14: 987-1001.
47. Mason, D. , Goneska, E., and Greider, C. W. (2003). Stem loop IV of Tetrahymena telomerase stimulates processivity in trans. Mol. Biol. Cell 23: 5606-5613
48. Chen, J.-L. and Greider, C.W. (2003). Determinants in mammalian telomerase RNA that mediate enzyme processivity and cross species incompatibility. Embo. J. 22: 304-314.
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49. Hackett, J. and Greider C.W. (2003). End resection initiates genomic instability in the absence of telomerase. Mol. Cell. Biol 23: 8450-8461.
50. Chen, J.-L., and Greider, C.W. (2003) Template boundary definition in mammalian telomerase. Genes Dev. 17: 2747-2752.
51. Qi, L., Strong, M., Karim, B. O., Huso, D. L., and Greider, C. W. (2003). Short telomeres and loss of ATM synergistically increase telomere dysfunction and suppress tumorigenesis. Cancer Research 63: 8188-8196.
52. Hao, L.-Y, and Greider, C.W. (2004). Genomic instability in both wildtype and telomerase-null MEFs. Chromsoma 113: 62-68.
53. Hao, L.-Y., and Greider, C. W. (2004). Phosphorylation of H2AX at short telomeres in T cells and fibroblasts. J. Biol. Chem. 43: 45148-45154.
54. Qi, L., Strong, M., Karim, B. O., Huso, D. L., and Greider, C. W. (2005). Telomere fusion to chromosome breaks reduced oncogenic translocations and tumor formation. Nat. Cell Biol. 7: 706-711.
55. Chen, J.-L., and Greider, C. W. (2005). Functional analysis of the pseudoknot structure in human telomerase RNA. Proc. Natl. Acad. Sci. U.S.A. 102: 8080-8085.
56. Armanios, M., Chen, J.-L., Chang, C. C., Brodsky, R. A., Hawkins, A., Griffen, C. A., J.R., E., Cohen, A. R., Chakravarti, A., Hamosh, A., and Greider, C. W. (2005). Haploinsufficiency of hTERT leads to anticipation in autosomal dominant dyskeratosis congenita. Proc. Natl. Acad. Sci. U.S.A. 102: 15960-15964.
57. Hao, L. Y., Armanios, M., Strong, M. A., Karim, B., Feldser, D. M., Huso, D., and Greider, C. W. (2005). Short Telomeres, even in the Presence of Telomerase, Limit Tissue Renewal Capacity. Cell 123: 1121-1131.
58. Feldser, D., Strong, M., and Greider, C.W. (2006) Ataxia telangiectasia mutated (Atm) is not required for telomerase-mediated elongation of short telomeres. Proc. Natl. Acad. Sci. U.S.A. 103: 2249-2251.
59. Frank, C. J., Hyde, M., and Greider, C. W. (2006). Regulation of telomere elongation by the cyclin-dependent kinase CDK1. Mol Cell, 24: 423-432.
60. Feldser, D., and Greider, C. W. (2007). Short telomeres limit tumor progression in vivo by inducing senescence. Cancer Cell: in press
61. Armanios, Y.M., Chen, J.-L., Cogan, J.D., Alder, J.K., Ingersoll, R.G., Markin, C., Lawson, W.E., Xiw, M., Vulto, I., Phillips, J.A., Lansdorp, P.M., Greider, C.W., and Loyd, J.E. (2007). Telomerase Mutations in Families with Idiopathic Pulmonary Fibrosis. New England Journal of Medicine in press.
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Reviews and Book Chapters:
1. Henderson, E., Larson, D., Melton, W., Shampay, J., Spangler, E., Greider, C.W., Ryan, T., and Blackburn, E.H. Structure, synthesis and regulation of telomeres. (1988) In: Cancer Cells 6: Eukaryotic DNA Replication. Cold Spring Harbor Laboratory. vol 6: 453–461.
2. Blackburn, E.H., Greider, C.W., Henderson, E., Lee, M., Shampay, J., and Shippen-Lentz, D. (1989). Recognition and elongation of telomeres by telomerase. Genome 31: 553–560.
3. Greider, C.W. (1990). Telomeres, telomerase and senescence. BioEssays 12: 363–369.
4. Greider, C.W. (1991). Chromosome first aid. Cell 67: 645–647.
5. Greider, C.W. (1991). Telomeres. Curr. Opin. Cell Biol. 3: 444–451.
6. Greider, C.W. (1992). Telomeres and telomerase in small eukaryotes. In: The eukaryotic genome: organization and regulation. P.M.A. Broda, S.G. Oliver and P.F.G Sims, eds. Society for General Microbiology Symposium 50. Cambridge University Press. pp 31-42.
7. Greider, C.W. (1992). Telomere chromatin and gene expression. Curr. Biol. 2: 62–64.
8. Greider, C.W. (1993). Telomerase and telomere length regulation: lessons from small eukaryotes to mammals. Cold Spring Harbor Symp. Quant. Biol. 58: 719-723.
9. Greider , C.W., Autexier, C., Avilion, A.A., Collins, K., Harrington, L.A., Mantell, L.L., Prowse, K.R., Smith, S.K., Allsopp, R.C., Counter, C.M., Vaziri, H., Bacchetti, S., Harley, C.B. (1993). Telomeres and telomerase: biochemistry and regulation in senescence and immortalization. In: The Chromosome. J.S. Hesslop-Harrison and R.B. Flavell, eds. Bios Scientific Publishers Ltd. pp. 115-125.
10. Harley, C.B., Kim, N.W., Prowse, K.R., Weinrich, S.L., Hirsch, K.S., West, M.D., Bacchetti, S., Hirte, H.W., Counter, C.M., Greider, C.W., Piatyszek, M.A., Wright, W.E., and Shay, J.W. (1994). Telomerase, cell immortality and cancer. Cold Spring Harbor Symp. Quant. Biol. 59: 307-315.
11. Greider, C.W. (1994). Mammalian telomere dynamics: healing, fragmentation, shortening and stabilization. Curr. Opin. Genet. and Dev., 4: 203-211.
12. Greider, C.W. (1995). Telomerase biochemistry and regulation. In: Telomeres. E.H. Blackburn and C.W. Greider, Eds. Cold Spring Harbor Laboratory Press. Cold Spring Harbor, New York. pp 35-68.
13. Greider, C. W., and Harley, C. B. (1996). Telomeres and telomerase in cell senescence and immortalization. In Cellular Aging and Cell Death, N. J. Holbrook, G. R. Martin, and R. A. Lockshin, eds. (New York, Wiley-Liss, Inc), pp. 123-138.
14. Greider, C.W. (1996). Telomere length regulation. Ann. Rev. Biochem. 65: 337-365.
15. Greider, C.W., and Blackburn, E.H. (1996). Telomeres, telomerase and cancer. Scientific American 274: 80-85.
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16. Greider, C.W., K. Collins, C. Autexier. (1996). Telomerases. In: DNA replication in Eukaryotic Cells. M.L. DePamphlis, Ed. Cold Spring Harbor Laboratory Press. Cold Spring Harbor New York. pp 619-638.
17. Autexier, C., and Greider, C.W. (1996). Telomerase and Cancer: revisiting the telomere hypothesis. Trends Biochem. 21: 387-391.
18. Greider, C.W. (1998). Telomerase activity, cell proliferation and cancer. Proc. Natl. Acad. Sci. U.S.A. 95: 90-92.
19. Greider, C.W. (1998). Telomeres and senescence: the history, the experiment, the future. Curr. Biol. 8: R178-R181.
20. Greider, C.W. (1999). Telomerase Activation: one step on the road to cancer?. Trends Genet. 15: 109-11.
21. Greider, C.W. (1999). Telomeres do D-loop-T-loop. Cell 97: 419-422.
22. Kass-Eisler, A., and Greider, C.W. (2000). Recombination and telomere-length maintenance. Trends Biochem. Sci. 25: 200-204.
23. Hemann, M. T., Hackett, J., IJpma, A., and Greider, C. W. (2000). Telomere length, telomere binding proteins and DNA damage signaling, Cold Spring Harbor Laboratory Symposium on Quantitative Biology LXV, 275-279.
24. Greider, C.W. 2001 Cellular responses to telomere shortening: cellular senescence as a tumor suppressor mechanism. Harvey Lect: 96: 33-50.
25. Hackett, J. A., and Greider, C. W. (2002). Balancing Instability: Dual roles for telomerase and telomere dysfunction in tumorigenesis, Oncogene, 21: 619-626.
26. Feldser, D. M., Hackett, J., and Greider, C. W. (2003). Telomere dysfunction and the initiation of genome instability. Nature Reviews Cancer 3: 623-627.
27. Greider, C.W., and Blackburn, E.H. (2004). Tracking Telomerase. Cell S11: S83-S86
28. Chen, J.L., and Greider, C.W. (2004). Telomerase RNA structure and function: implications for dyskeratosis congentia Trends Biochem Sci. 29: 183-192.
29. Chen, J.L., and Greider, C.W. (2004). An emerging consensus for telomerase RNA structure. Proc. Natl. Acad. Sci. U.S.A. 101: 14683-14684.
30. Armanios, M., and Greider, C.W. (2005). Telomerase and Cancer Stem Cells. Cold Spring Harbor Laboratory Symposium on Quantitative Biology. Vol. LXX 205-208.
31. Chen, J.-L., and Greider, C. W. (2006). Telomerase Biochemistry and Biogenesis. In Telomeres, T. DeLange, E. H. Blackburn, and V. Lundblad, Cold Spring Harbor, NY, Cold Spring Harbor Press 49-73.
32. Blackburn, E. H., Greider, C. W., and Szostak, J. W. (2006). Telomeres and telomerase: the
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path from maize, Tetrahymena and yeast to human cancer and aging. Nat. Med. 12, 1133-1138.
33. Greider, C.W. (2007) Telomerase RNA Levels Limit the Telomere Length Equilibrium. Cold Spring Harbor Laboratory Symposium on Quantitative Biology. Vol. LXXI 225-229.

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    Faculty position at Johns Hopkins University