Carol Widney Greider, Ph.D.

Headshot of Carol Widney Greider
  • Daniel Nathans Professor and Director, Department of Molecular Biology and Genetics
  • Professor of Molecular Biology and Genetics

Research Interests

Telomeres and Telomerase in Cancer and age related degenerative disease more


Carol Greider, Ph.D. received her bachelor’s degree from the University of California at Santa Barbara in 1983 and a Ph.D. in 1987 from the University of California at Berkeley. In 1984, working together with Dr. Elizabeth Blackburn, she discovered telomerase, an enzyme that maintains telomeres, or chromosome ends. In 1988, Dr. Greider went to Cold Spring Harbor Laboratory where, as an independent Cold Spring Harbor Fellow, she cloned and characterized the RNA component of telomerase. In 1990, Dr. Greider was appointed as an assistant investigator at Cold Spring Harbor Laboratory, followed later by appointment to Investigator in 1994. She expanded the focus of her telomere research to include the role of telomere length in cellular senescence, cell death and in cancer.  In 1997, Dr. Greider moved her laboratory to the Department of Molecular Biology and Genetics at The Johns Hopkins University School of Medicine. In 2003 she was appointed as the Daniel Nathans Professor and Director of the Department of Molecular Biology and Genetics. At Johns Hopkins University, Dr. Greider’s group continued to study the biochemistry of telomerase and determined the secondary structure of the human telomerase RNA. In addition she characterized the loss of telomere function in mice, which allowed an understanding of humans short telomere diseases such as bone marrow and other stem cell failure diseases. Dr. Greider shared the Nobel Prize in Physiology or Medicine in 2009 with Drs. Elizabeth Blackburn and Jack Szostak for their work on telomeres and telomerase. Dr. Greider currently directs a group of eight scientists studying both the role of short telomeres in age-related disease and cancer as well as the regulatory mechanism that maintain telomere length. more


  • Daniel Nathans Professor and Director, Department of Molecular Biology and Genetics
  • Bloomberg Distinguished Professor
  • Professor of Molecular Biology and Genetics
  • Professor of Oncology

Departments / Divisions

Centers & Institutes



  • Ph.D.; University of California (Berkeley) (California) (1987)
  • B.A.; University of California (Santa Barbara) (California) (1983)

Additional Training

  • Cold Spring Harbor Laboratory, Cold Spring Harbor, NY, 1990

Research & Publications

Research Summary

Telomeres protect chromosome ends from being recognized as DNA damage and chromosomal rearrangements. Conventional replication leads to telomere shortening, but telomere length is maintained by the enzyme telomerase that synthesizes telomere sequences de novo onto chromosome ends. Telomerase is specialized reverse transcriptase, requiring both a catalytic protein and an essential RNA component. In the absence of telomerase, telomeres shorten progressively as cells divide, and telomere function is lost. For this reason, telomerase is required for cells that undergo many rounds of divisions, especially tumor cells and some stem cells. My lab is focused understanding telomerase and cellular and organismal consequences of telomere dysfunction. We use biochemistry, yeast and mice to examine telomere function. We generated telomerase null mice that are viable and show progressive telomere shortening for up to six generations. In the later generations, when telomeres are short, cells die via apoptosis or senescence. Crosses of these telomerase null mice to other tumor prone mice show that tumor formation can be greatly reduced by short telomeres. We also are using our telomerase null mice to explore the essential role of telomerase stem cell viability. Telomerase mutations cause telomere syndromes in humans due to loss of tissue renewal capacity. Uncovering the mechanisms that regulate the telomere length equilibrium will allow us to understand these age-related degenerative diseases, and suggest avenues for treatment in the future 


Lab Website: Greider Lab

Selected Publications

View all on PubMed

Publications: Peer-reviewed Original Science Research

1.     Greider, C.W., and Blackburn, E.H.  Identification of a specific telomere terminal transferase activity in Tetrahymena extracts. Cell  (1985) 43: 405–413.

2.     Greider, C.W. and Blackburn, E.H.  The telomere terminal transferase of Tetrahymena is a ribonucleoprotein enzyme with two kinds of primer specificity. Cell (1987) 51: 887–898.

3.     Greider, C.W., and Blackburn, E.H.  A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis. Nature (1989) 337: 331–337.

4.     Harley, C.B., Futcher, A.B., and Greider, C.W.  Telomeres shorten during ageing of human fibroblasts. Nature  (1990) 345: 458–460.

5.     Harrington, L.A., and Greider, C.W.  Telomerase primer specificity and chromosome healing. Nature (1991) 353: 451–454.

6.     Greider, C.W.  Telomerase is processive. Mol. Cell Biol.  (1991) 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.  Telomere length predicts the replicative capacity of human fibroblasts. Proc. Natl. Acad. Sci. U.S.A. (1992) 89: 10114–10118.

8.     Levy, M.Z., Allsopp, R.C., Futcher, A.B., Greider, C.W., and Harley, C.B.  Telomere end replication problem and cell aging. J. Mol. Biol. (1992) 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.  Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity. EMBO J. (1992) 11: 1921–1929.

10.  Avilion, A.A., Harrington, L.A., and Greider, C.W.  Tetrahymena telomerase RNA levels increase during macronuclear development. Dev. Genet. (1992) 13: 80–86.

11.  Collins, K. and Greider, C.W.  Tetrahymena telomerase catalyzes nucleolytic cleavage and nonprocessive elongation. Genes Dev. (1993) 7: 1364-1376.

12.  Prowse, K.R., Avilion, A.A., and Greider, C.W.  Identification of a nonprocessive telomerase activity from mouse cells. Proc. Natl. Acad. Sci. U.S.A. (1993) 90: 1493-1497.

13.  Mantell, L.L., and Greider, C.W.  Telomerase activity in germline and embryonic cells of Xenopus. EMBO J. (1994) 13: 3211-3217.

14.  Autexier, C., and Greider, C.W.  Functional reconstitution of wild-type and mutant Tetrahymena telomerase. Genes Dev. (1994) 8: 563-575.

15.  Collins, K., and Greider, C.W.  Utilization of ribonucleotides and RNA primers by Tetrahymena telomerase. EMBO J. (1995) 14: 5422-5432.

16.  Autexier, C., and Greider, C.W.  Boundary elements of the Tetrahymena telomerase RNA template and alignment domains. Genes & Dev. (1995) 9: 2227-2239.

17.  Blasco, M.A., Funk, W., Villeponteau, B., and Greider, C.W.  Functional characterization and developmental regulation of the mouse telomerase RNA. Science (1995) 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.  The RNA component of human telomerase. Science (1995) 269: 1236-1241.

19.  Collins, K., Kobayashi, R., and Greider, C.W.  Purification of Tetrahymena telomerase and cloning of genes encoding the two protein components of the enzyme. Cell (1995) 81: 677-686.

20.  Prowse, K.R., and Greider, C.W.  Developmental and tissue specific regulation of mouse telomerase and telomere length. Proc. Natl. Acad. Sci. U.S.A. (1995) 92: 4818-4822.

21.  Harrington, L.A., Hull, C., Crittenden, J., and Greider, C.W.  Gel shift and UV cross-linking analysis of Tetrahymena telomerase. J. Biol. Chem. (1995) 270: 8893-8901.

22.  Blasco, M., Rizen, M., Greider, C.W., and Hanahan, D.  Differential regulation of telomerase activity and telomerase RNA during multi-stage tumorigenesis. Nat. Genet. (1996) 12: 200-204.

23.  Avilion, A.A., Piatyszek, M. A., Gupta, J., Shay, J.W., Bacchetti, S., and Greider, C.W.  Human telomerase RNA and telomerase activity in immortal cell lines and tumor tissues. Cancer Res. (1996) 56: 645-650.

24.  Buchkovich, K.J., and Greider, C.W.  Telomerase regulation during entry into the cell cycle in normal human T cells. Mol. Biol. Cell (1996) 7: 1443-1454.

25.  Autexier, C., Pruzan, R., Funk, W., and Greider, C.W.  Reconstitution of human telomerase activity and identification of a minimal functional region of the human telomerase RNA. EMBO J. (1996) 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.  Telomere shortening and tumor formation by mouse cells lacking telomerase RNA. Cell (1997) 91: 25-34.

27.  Hinkley, C.S., Blasco, M.A., Funk, W.D., Feng, J., Villeponteau, B., Greider, C.W., and Herr, W.  The mouse telomerase RNA 5’-end lies just upstream of the telomerase template sequence. Nucleic Acids Res. (1998) 26: 532-536.

28.  Le, S., Zhu, J.J., Anthony, D.C., Greider, C.W., and Black, P.M.  Telomerase activity in human gliomas. Neurosurgery (1998) 42: 1120-1125.

29.  Autexier, C.A., and Greider, C.W.  Mutational analysis of the Tetrahymena telomerase RNA: Identification of residues affecting telomerase activity in vitro. Nucleic Acids Res. (1998) 26: 787-795.

30.  Lee, H.-W., Blasco, M.A., Gottlieb, G.J., Horner, J.W., Greider, C.W., and DePinho, R.A.  Essential role of mouse telomerase in highly proliferative organs. Nature (1998) 392: 569-574.

31.  Rudolph, K.L., Chang, S., Lee, H.-W., Blasco, M., Gottlieb, G., Greider, C.W., and DePinho, R.A.  Longevity, stress response, and cancer in aging telomerase deficient mice. Cell (1999) 96: 701-712.

32.  Le, S., Moore, J.K., Haber, J.E., and Greider, C.W.  RAD51 and RAD50 define two pathways that collaborate to maintain telomeres in the absence of telomerase. Genetics (1999) 152: 143-152.

33.  Greenberg, R., Chin, L., Femino, A., Lee, K.-H., Gottlieb, G., Singer, R., Greider, C.W., and DePinho, R.A.  Short telomeres impair tumorigenesis in the INK4aD2/3-/- mTR-/- cancer-prone mouse. Cell (1999) 97: 515-525.

34.  Chin, L., Artandi, S., Shen, Q., Tam, S., Lee, S.-L., Gottlieb, G., Greider, C.W., and DePinho, R.A.  p53 deficiency rescues the adverse effects of telomere loss in vivo and cooperates with telomere dysfunction to accelerate carcinogenesis. Cell (1999) 97: 527-538.

35.  Hemann, M.T., and Greider, C.W.  G-strand overhangs on telomeres in telomerase deficient mouse cells. Nucleic Acids Res. (1999) 27: 3964-3969.

36.  Le, S., Sternglanz, R., and Greider, C.W.  Identification of two RNA binding proteins associated with human telomerase RNA. Mol. Biol. Cell. (2000) 11: 999-1010.

37.  Chen, J.-L., Blasco, M., and Greider, C.W.  A Secondary structure of vertebrate telomerase RNA. Cell (2000) 100: 503-514.

38.  Hemann, M.T., and Greider, C.W.  Wild derived inbred mouse strains have short telomeres, Nucleic Acids Res. (2000) 28: 4474-4478.

39.  Chen, Q., IJpma, A., and Greider, C.W.  Two survivor pathways that allow growth in the absence of telomerase are generated by distinct telomere recombination events. Mol. Cell. Biol. (2001) 21: 1819-1827.

40.  Hemann, M.T., Rudolph, L., Strong, M., DePinho, R.A., Chin, L., and Greider, C.W.  Telomere dysfunction triggers developmentally regulated germ cell apoptosis.  Mol. Biol. Cell. (2001) 12: 2023-2030.

41.  Hackett, J., Feldser, D.M., and Greider, C.W.  Telomere dysfunction increases mutation rate and genomic instability. Cell (2001) 106: 275-286.

42.  Hemann, M.T., Strong, M., Hao, L.-Y., and Greider, C.W.  The shortest telomere, not average telomere length, is critical for cell viability and chromosome stability. Cell (2001) 107: 66-77.

43.  Mason, D., Autexier, C., and Greider, C.W.  Tetrahymena proteins p80 and p95 are not core telomerase components. Proc. Natl. Acad. Sci. U.S.A. (2001) 98: 12368-12373.

44.  Chen, J.-L., Opperman, K.K., and Greider, C.W.  A critical stem loop structure in the CR4-CR5 domain of mammalian telomerase RNA. Nucleic Acids Res. (2002) 30: 592-597.

45.  Hathcock, K.S., Hemann, M.T., Opperman, K.K., Strong, S.A., Greider, C.W., and Hodes, R.J.  Haploinsufficiency of mTR results in defects in telomere elongation.  Proc. Natl. Acad. Sci. U.S.A. (2002) 99: 3591-3596.

46.  IJpma, A., and Greider, C.W.  Short telomeres induce a DNA damage response in S. cerevisiae. Mol. Biol. Cell(2003) 14: 987-1001.

47.  Mason, D., Goneska, E., and Greider, C.W.   Stem loop IV of Tetrahymena telomerase stimulates processivity in trans. Mol. Biol. Cell(2003) 23: 5606-5613.

48.  Chen, J.-L. and Greider, C.W.  Determinants in mammalian telomerase RNA that mediate enzyme processivity and cross species incompatibility. Embo. J. (2003) 22: 304-314.

49.  Hackett, J. and Greider, C.W.  End resection initiates genomic instability in the absence of telomerase. Mol. Cell. Biol (2003) 23: 8450-8461.

50.  Chen, J.-L., and Greider, C.W.  Template boundary definition in mammalian telomerase.Genes Dev. (2003) 17: 2747-2752.

51.  Qi, L., Strong, M., Karim, B.O., Huso, D.L., and Greider, C.W.  Short telomeres and loss of ATM synergistically increase telomere dysfunction and suppress tumorigenesis. Cancer Research (2003) 63: 8188-8196.

52.  Hao, L.-Y, and Greider, C.W.  Genomic instability in both wildtype and telomerase-null MEFs. Chromsoma (2004) 113: 62-68.

53.  Hao, L.-Y., and Greider, C.W.  Phosphorylation of H2AX at short telomeres in T cells and fibroblasts. J. Biol. Chem. (2004) 43: 45148-45154.

54.  Qi, L., Strong, M., Karim, B.O., Huso, D.L., and Greider, C.W.  Telomere fusion to chromosome breaks reduced oncogenic translocations and tumor formation. Nat. Cell Biol. (2005) 7: 706-711.

55.  Chen, J.-L., and Greider, C.W.  Functional analysis of the pseudoknot structure in human telomerase RNA. Proc. Natl. Acad. Sci. U.S.A. (2005) 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.  Haploinsufficiency of hTERT leads to anticipation in autosomal dominant dyskeratosis congenita. Proc. Natl. Acad. Sci. U.S.A. (2005) 102: 15960-15964.

57.  Hao, L.Y., Armanios, M., Strong, M.A., Karim, B., Feldser, D.M., Huso, D., and Greider, C.W.  Short Telomeres, even in the Presence of Telomerase, Limit Tissue Renewal Capacity. Cell (2005) 123: 1121-1131.

58.  Feldser, D., Strong, M., and Greider, C.W.  Ataxia telangiectasia mutated (Atm) is not required for telomerase-mediated elongation of short telomeres. Proc. Natl. Acad. Sci. U.S.A. (2006) 103: 2249-2251.

59.  Frank, C.J., Hyde, M., and Greider, C.W.  Regulation of telomere elongation by the cyclin-dependent kinase CDK1. Mol Cell  (2006) 24: 423-432.

60.  Feldser, D., and Greider, C.W.   Short telomeres limit tumor progression in vivo by inducing senescence. Cancer Cell (2007) 11: 461-469.

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.  Telomerase Mutations in Families with Idiopathic Pulmonary Fibrosis. The New England Journal of Medicine (2007) 356: 1317-1326.

62.  Morrish, T.A., Greider, C.W.   Short Telomeres Initiate Telomere Recombination in Primary and Tumor Cells. PLoS Genet. (2009) 5 (1): e1000357.

63.  Ma, Y., and Greider, C.W.  Kinase-Independent Functions of TELI in Telomere Maintenance. Mol Cell Biol. (2009) 29:  5193-5202.

64.  Armanios, M., Alder, J.K., Parry, E.M., Karim, B., Strong, M.A., and Greider, C.W.  Short telomeres are sufficient to cause the degenerative defects associated with aging. The American Journal of Human Genetics (2009) 85: 823-832.

65.  Vidal-Cardenas, S.L., and Greider, C.W.  Comparing effects of mTR and mTERT deletion on gene express and DNA damage response: a critical examination of telomere length maintenance-independent roles of telomerase.  Nucleic Acids Res. (2010) 38: 60-71.

66.  Tom, H.I., and Greider, C.W.  A Sequence-Dependent Exonuclease Activity From Tetrahymena thermophila; BMC Biochem. (2010) 11:45.

67.  Strong, M.A.,Vidal-Cardenas, S.L., Karim, B., Yu, H., Guo, N., Greider, C.W.  Phenotypes in mTERT+/- and mTERT-/- Mice are Due to Short Telomeres, Not Telomere-Independent Functions of TERT. Mol Cell Biol. (2011) 31: 2369-2379.

68.  Bender H.S., Murchison E.P., Pickett H.A., Deakin J.E., Strong M.A., Conlan C., McMillan

D.A., Neumann, A.A., Greider, C.W., Hannon, G.J., Reddel, R.R., Graves, J.A.  Extreme telomere length dimorphism in the Tasmanian devil and related marsupials suggests parental control of telomere length. PLoS One. (2012) 7(9): e46195.

69. Kaizer, H., Connelly, C.J., Bettridge, K., Viggiani, C., and Greider, C.W. Regulation of Telomere Length Requires a Conserved N-Terminal Domain of Rif2 in Saccharomyces cerevisiae. Genetics (2015) 201:573-586.

70. Lee, S. S., Bohrson, C., Pike, A.M., Wheelan, S.J.,  and Greider, C.W. ATM kinase is required for telomere elongation in mouse and human cells. Cell Reports. (2015) in press.

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.  In:  Cancer Cells 6:  Eukaryotic DNA Replication. Cold Spring Harbor Laboratory. Vol. (1988) 6:  453–461.

2.     Blackburn, E.H., Greider, C.W., Henderson, E., Lee, M., Shampay, J., and Shippen-Lentz, D.  Recognition and elongation of telomeres by telomerase. Genome (1989) 31: 553–560.

3.     Greider, C.W.  Telomeres, telomerase and senescence. BioEssays  (1990) 12: 363–369.

4.     Greider, C.W.  Chromosome first aid. Cell (1991) 67: 645–647.

5.     Greider, C.W.  Telomeres. Curr. Opin. Cell Biol. (1991) 3: 444–451.

6.     Greider, C.W. 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. (1992) pp. 31-42.

7.     Greider, C.W.  Telomere chromatin and gene expression. Curr. Biol. (1992) 2:  62–64.

8.     Greider, C.W.  Telomerase and telomere length regulation:  lessons from small eukaryotes to mammals. Cold Spring Harbor Symp. Quant. Biol. (1993) 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.  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. (1993) 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.  Telomerase, cell immortality and cancer. Cold Spring Harbor Symp. Quant. Biol. (1994) 59: 307-315.

11.  Greider, C.W.  Mammalian telomere dynamics:  healing, fragmentation, shortening and stabilization. Curr. Opin. Genet. and Dev. (1994) 4: 203-211.

12.  Greider, C.W.  Telomerase biochemistry and regulation. In:  Telomeres. E.H. Blackburn and C.W. Greider, Eds. Cold Spring Harbor Laboratory Press. Cold Spring Harbor, New York (1995) pp. 35-68.

13.  Greider, C.W., and Harley, C.B.  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. (1996) pp. 123-138.

14.  Greider, C.W.  Telomere length regulation. Ann. Rev. Biochem. (1996) 65: 337-365.

15.  Greider, C.W., and Blackburn, E.H.  Telomeres, telomerase and cancer.  Scientific American (1996) 274:  92-97.

16.  Greider, C.W., K. Collins, C. Autexier.  Telomerases. In:  DNA replication in Eukaryotic Cells. M.L. DePamphlis, Ed. Cold Spring Harbor Laboratory Press. Cold Spring Harbor New York (1996) pp. 619-638.

17.  Autexier, C., and Greider, C.W.  Telomerase and Cancer:  revisiting the telomere hypothesis. Trends Biochem. (1996) 21:  387-391.

18.  Greider, C.W.  Telomerase activity, cell proliferation and cancer. Proc. Natl. Acad. Sci. U.S.A. (1998) 95:  90-92.

19.  Greider, C.W.  Telomeres and senescence:  the history, the experiment, the future. Curr. Biol. (1998) 8:  R178-R181.

20.  Greider, C.W.  Telomerase Activation:  one step on the road to cancer? Trends Genet. (1999) 15: 109-112.

21.  Greider, C.W.  Telomeres do D-loop-T-loop. Cell (1999) 97: 419-422.

22.  Kass-Eisler, A., and Greider, C.W.  Recombination and telomere-length maintenance. Trends Biochem. Sci. (2000) 25: 200-204.

23.  Hemann, M.T., Hackett, J., IJpma, A., and Greider, C.W.  Telomere length, telomere binding proteins and DNA damage signaling, Cold Spring Harbor Laboratory Symposium on Quantitative Biology LXV (2000) 275-279.

24.  Greider, C.W.  Cellular responses to telomere shortening: cellular senescence as a tumor suppressor mechanism. Harvey Lect: (2001) 96: 33-50.

25.  Hackett, J.A., and Greider, C.W.  Balancing Instability: Dual roles for telomerase and telomere dysfunction in tumorigenesis, Oncogene (2002) 21: 619-626.

26.  Feldser, D.M., Hackett, J., and Greider, C.W.  Telomere dysfunction and the initiation of genome instability. Nature Reviews Cancer (2003) 3: 623-627.

27.  Greider, C.W., and Blackburn, E.H.  Tracking Telomerase. Cell (2004) S11: S83-S86

28.  Chen, J.L., and Greider, C.W.  Telomerase RNA structure and function: implications for dyskeratosis congentia Trends Biochem Sci. (2004) 29: 183-192.

29.  Chen, J.L., and Greider, C.W.  An emerging consensus for telomerase RNA structure.  Proc. Natl. Acad. Sci. U.S.A. (2004) 101: 14683-14684.

30.  Armanios, M., and Greider, C.W.  Telomerase and Cancer Stem Cells. Cold Spring Harbor Laboratory Symposium on Quantitative Biology. Vol. LXX (2005) 205-208.

31.  Chen, J.-L., and Greider, C.W.  Telomerase Biochemistry and Biogenesis. In Telomeres, T. DeLange, E.H. Blackburn, and V. Lundblad, Cold Spring Harbor, NY, Cold Spring Harbor Press  (2006) 49-73.

32.  Blackburn, E.H., Greider, C.W., and Szostak, J.W.  Telomeres and telomerase: the path from maize, Tetrahymena and yeast to human cancer and aging. Nat. Med. (2006) 12: 1133-1138.

33.  Greider, C.W.  Telomerase RNA Levels Limit the Telomere Length Equilibrium. Cold Spring Harbor Laboratory Symposium on Quantitative Biology. Vol. LXXI (2007) 225-229.

34.  Greider, C.W.  Telomerase Discovery: The Excitement of Putting Together Pieces of the Puzzle (Nobel Lecture). Angew Chem Int Ed Engl. (2010) 49: 7422-7439.

35.  Greider, C.W.  Els telòmers i la telomerasa: una breu història sobre els telòmers i el descobriment del seu paper en les malalties que afecten els humans (Telomeres and telomerase:  a short history of telomeres and uncovering their role in human disease).  Revista de la Reial Academia de Medicina de Catalunya. (2011) 26, 142.

36.  Greider, C.W.  Molecular Biology. Wnt regulates TERT--putting the horse before the cart. Science (2012) 336 (6088): 1519-1520.

37.  Armanios, M., and Greider, C.W. Treating Myeloproliferation--On Target or Off? The New England Journal of Medicine (2015) 373(10): 965-966.



1.       Craig, N.L., Cohen-Fix, O., Green, R., Greider, C.W., Storz, G., and Wolberger, C.  Molecular Biology: Principles of Genome Function (2010) (Oxford, Oxford University Press).

2.       Craig, N.L., Cohen-Fix, O., Green, R., Greider, C.W., Storz, G., and Wolberger, C.  Molecular Biology: Principles of Genome Function, Second Edition (2014) (Oxford, Oxford University Press).




Contact for Research Inquiries

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Fax: 410-955-0831

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Academic Affiliations & Courses

Graduate Program Affiliation

Biochemistry, Cellular and Molecular Biology Graduate Program

Cellular and Molecular Medicine

Human Genetics and Molecular Biology

Activities & Honors


  • Fellow, American Academy of Arts and Sciences, 2003
  • Fellow, American Academy of Microbiology, 2004
  • Cornelius Rhoads Award, American Association for Cancer Research, 1996
  • Gertrude Elion Cancer Research Award, American Association for Cancer Research, 1994
  • Fellow, American Association for the Advancement of Science, 2003
  • Schering-Plough Scientific Achievement Award, American Society for Biochemistry and Molecular Biology, 1997
  • Glenn Foundation Award, American Society for Cell Biology, 1995
  • Lewis S. Rosenstiel Award for Distinguished Work in Basic Medical Research, Brandeis University, 1999
  • Louisa Gross Horwitz Prize, Columbia University, 2007
  • Albert Lasker Award for Basic Medical Research, Lasker Foundation, 2006
  • Katharine Berkan Judd Award, Memorial Sloan Kettering Cancer Center, 2008
  • Richard Lounsbery Award, National Academy of Science, 2003
  • Paul Ehrlich and Ludwig Darmstaedter Prize, Paul Ehrlich Foundation, 2009
  • Nobel Prize in Physiology or Medicine, Royal Swedish Academy of Sciences, 2009
  • Pearl Meister Greengard Prize, The Rockefeller University, 2009
  • Regents Scholarship, University of California, 1981
  • Dickson Prize in Medicine, University of Pittsburgh, 2007
  • Wiley Prize in Biomedical Sciences, Wiley Foundation, 2006
  • Lila and Murray Gruber Memorial Cancer Research Award, 2006
  • Passano Foundation Award, 1999
  • Gairdner Foundation Award, 1998
  • Senior Scholar, Ellison Medical Foundation , 1998
  • Allied Signal Outstanding Project Award, 1992
  • Pew Scholar in the Biomedical Sciences, 1990


  • National Academy of Sciences, 2003

Professional Activities

  • Appointee, President's Committee on the National Medal of Science, 2012 - 2014

Videos & Media

Lectures and Presentations

  • Harvey Society Lecture
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