Selected Publications

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

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.

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

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

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.

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.

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.

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.

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

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.

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

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.

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

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

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.

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.

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.

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

Morrish, T.A., and Greider, C.W. (2009). Short telomeres initiate telomere recombination in primary and tumor cells. PLoS Genetics 5: e1000357.

Ma, Y., and Greider, C.W. (2009). Kinase-independent functions of TEL1 in telomere maintenance. Mol. Cell. Biol. 29: 5193-5202.

Vidal-Cardenas, S.L., and Greider, C.W. (2009). Comparing effects of mTR and mTERT deletion on gene expression and DNA damage response: a critical examination of telomere length maintenance-independent roles of telomerase. Nucleic Acids Res. 38: 60-71.

Armanios, M., Alder, J.K., Parry, E.M., Karim, B., Strong, M.A., and Greider, C.W. (2009). Short telomeres are sufficient to cause the degenerative defects associated with aging. Am J Hum Genet. 85: 823-832.

Greider, C.W. (2010). Telomerase discovery: the excitement of putting together pieces of the puzzle (Nobel lecture). Angew Chem Int Ed Engl 49: 7422-7439.

Tom, HI., and Greider, C.W. (2010). A sequence-dependent exonuclease activity from Tetrahymena thermophila. BMC Biochem 11: 45.

Strong MA, Vidal-Cardenas SL, Karim B, Yu H, Guo N, Greider CW (2011). Phenotypes in mTERT+/- and mTERT-/- mice are due to short telomeres, not telomere-independent functions of telomerase reverse transcriptase. Mol. Cell Biol. Jun;31(12):2369-79. Epub 2011 Apr 4.

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

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., and Graves, J.A. (2012). Extreme telomere length dimorphism in the Tasmanian devil and related marsupials suggests parental control of telomere length. PLoS One 7(9): e46195. Epub 2012 Sep 25.

Kaizer H, Connelly CJ, Bettridge K, Viggiani C, and Greider CW. (2015). Regulation of Telomere Length Requires a Conserved N-Terminal Domain of Rif2 in Saccharomyces cerevisiae. Genetics Oct;201(2):573-86. Epub 2015 Aug 20.

Armanios M, and Greider CW. (2015). Treating myeloproliferation - on target or off? N Engl J Med Sep 3;373(10): 956-6.

Lee SS, Bohrson C, Pike AM, Wheelan SJ, and Greider CW. (2015). ATM kinase is required for telomere elongation in mouse and human cells. Cell Reports 13(8): 1623-32. Epub 2015 Nov 12.

Wang S, Pike AM, Lee SS, Strong MA, Connelly CJ, Greider CW. (2017). BRD4 inhibitors block telomere elongation. Nucleic Acids Res. Aug 21;45(14):8403-8410.

Alder JK, Hanumanthu VS, Strong MA, DeZern AE, Stanley SE, Takemoto CM, Danilova L, Applegate CD, Bolton SG, Mohr DW, Brodsky RA, Casella JF, Greider CW, Jackson JB, Armanios M. (2018). Diagnostic utility of telomere length testing in a hospital-based setting. Proc. Natl. Acad. Sci. U.S.A. Mar 6;115(10):E2358-E2365. Epub 2018 Feb 20.

Pike AM, Strong MA, Ouyang JPT, Greider CW. (2019). TIN2 functions with TPP1/POT1 to stimulate telomerase processivity. Mol. Cell Biol. 39(21). Epub 2019 Oct 11.

Keener R, Connelly CJ, Greider CW. (2019). Tel1 activation by the MRX complex is sufficient for telomere length regulation but not for the DNA damage response in Saccharomyces cerevisiae. Genetics. Epub 2019 Oct 23.

Shubin CB and Greider CW. (2020). The role of Rif1 in telomere length regulation is separable from its role in origin firing. Elife. Epub 2020 Jun 29.

Shubin CB, Mayangsari R, Swett AD, Greider CW. (2021). Rif1 regulates telomere length through conserved HEAT repeats. NAR. Epub 2021 Mar 27.


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

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

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

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

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

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.

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

Greider, C.W. (2007) Telomerase RNA levels limit the telomere length equilibrium. Cold Spring Harbor Laboratory Symposium on Quantitative Biology. Vol. LXXI 225-229.

Greider C.W. (2016). Regulating telomere length from the inside out: the replication fork model. Genes Dev. 30(13):1483-91.