Birth Date: 15 April 1961
Place of Birth: San Diego, California , USA
Nationality: American
Occupation/Field of Study American molecular biologist, co-recipient of a Nobel Prize with Elizabeth Blackburn and Jack Szostak for the discovery of how chromosomes are protected by telomeres and the enzyme telomerase


KEYWORDS: Nobel Prize, USA, Carol W. Greider, Marine Ecology, Telomeres



Carol Greider is an American scientist and holder of a Degree in Marine Ecology and a PhD in Molecular Science who has worked side by side with Elizabeth Blackburn, with whom she discovered telomeres (enzymes at the end of chromosomes which contain RNA). This discovery, which led her to share a Nobel Prize with Blackburn, enhanced the study of ageing, cancer and related diseases.



Greider was born in California in 1961, the second child (she has an older brother Mark) of her parents, who were both postdoctoral researchers; her father in Physics and her mother in Mycology. She lost her mother at the age of six. In 1971, the family moved to Heidelberg, Germany for a year. Greider studied Marine Ecology at Santa Barbara, where she spent much of her time in the laboratory, and then completed her junior year at the University of Göttingen, working on chromosomes2. Greider didn’t score well on her graduate exams because of her dyslexia, but she was still accepted to UC Berkeley, where she met Elizabeth Blackburn2 and, in 1984, started working on telomeres in her laboratory. After obtaining a PhD in Molecular Biology at the end of 1987, she began a postdoctoral fellowship at Cold Spring Harbor Laboratory.

Greider married Nathaniel Comfort in 1993, a science historian with whom she has two children, son Charles and daughter Gwendolyn. Greider and her husband Nathaniel held positions at universities in neighbouring cities since 1997 up until 2003 when Comfort also joined the Johns Hopkins University School of Medicine2,3. The couple is now divorced. In 1997, she started working in Tom Kelly’s department of Molecular Biology and Genetics at Johns Hopkins University School of Medicine in Baltimore.

Carol Greider (img courtesy Newswise)

In 1996, President Clinton appointed Greider to the U.S. Congress’s National Bioethics Advisory Commission4, which dealt with issues such as cloning and stem cell research. The commission was succeeded by the President’s Council on Bioethics in 2001, at which time Elizabeth Blackburn was also appointed to it, serving until 2004 when she was ousted to much scientific outcry after her continued support of stem cell research4. Greider has expressed that she feels obliged as a research scientist to engage in the discussion of controversial subjects, so she can give the point of view of a specialist5.

In 2003, Greider was named Daniel Nathans Professor and Department Director for Molecular Biology and Genetics at the Johns Hopkins University School of Medicine. She was appointed Bloomberg Distinguished Professor at the School’s Department of Biology in 2014.



Greider was Blackburn’s graduate student when the two of them made the discovery of the telomeres, enzymes at the end of the chromosomes that contain RNA. As the telomeres shorten, cells stop dividing. This is a crucial factor when it comes to ageing, cancer and related diseases. Just like Blackburn, Greider continues to work on this field of research6.



Having two kids and a full time job in the lab is a challenge, but having Charles and Gwendolyn is the best thing that has ever happened to me. My lab knows that I am a mom first, and the flexibility that academic science provides makes having a career and a family possible. […] The main thing is to find the time to get things done, it is not the hours at work but the overall productivity that counts. Having flexibility takes a huge amount of pressure off.

“Carol W. Greider – Biographical” (2009),, Nobel Media AB,  <> (last accessed 13 Jun. 2016).


If we’re going to have 50% of the people that may have creative ideas not really be able to engage, than that certainly takes away a lot of the possibilities for discoveries. […] Science is to the detriment because of that. […] That’s a larger issue  […] I don’t think science is unique in this area.

“Scientific Discovery with Carol Greider – Conversations with History”(6 May 2014), Youtube, uploaded by University of California Television (UCTV), <> (last accessed 13 Jun. 2016).


When I was at Berkeley in graduate school, 50% of us were women, but that has not tracked along  – as you would expect […], it’s a not a pipeline issue, [according to the report by the Association of University Women], it’s a leaky pipeline: although there are 50% of women coming in, […] at each stage more and more women decide not to go on in the pipeline, so they’re coming out of the pipeline.

“Scientific Discovery with Carol Greider  – Conversations with History” (6 May 2014), Youtube, uploaded by University of California Television (UCTV), <> (last accessed 13 Jun. 2016).



1994: Gertrude Elion Cancer Research Award (AACR)
1995: Glenn Foundation Award from American Society for Cell Biology (ASCB)
1996: Cornelius Rhoads Award (AACR)
1997: Schering-Plough Scientific Achievement Award, American Society for Biochemistry and Molecular Biology
1998: Canada Gairdner Foundation International Award, with Elizabeth Blackburn
1998: The Lewis S. Rosenstiel Award with Elizabeth Blackburn “for their outstanding work on the maintenance of telomeres”7
1999: Member of the American Society for Cell Biology (ASCB)
1999: The Passano Foundation Award with Elizabeth Blackburn “for determining the molecular nature of telomeres (chromosome ends) and discovering the enzyme telomerase, which carries out the synthesis of the telomeres”8
2000: Academy of Achievement Golden Plate Award, with Elizabeth Blackburn
2003: Fellow of the American Academy of Arts and Sciences and of the American Association for the Advancement of Science
2003: Member of the National Academy of Sciences
2003: The Richard Lounsbery Award from the National Academy of Sciences “for her pioneering biochemical and genetic studies of telomerase, the enzyme that maintains the ends of chromosomes in eukaryotic cell”9
2004: Member of the American Society for Biochemistry and Molecular Biology
2006: Albert Lasker Award for Basic Medical Research, shared with Elizabeth Blackburn and Jack Szostak, for predicting and discovering telomerase
2006: Wiley Prize in Biomedical Sciences, shared with Elizabeth Blackburn
2006: Lila Gruber Cancer Research Award for outstanding contributions in the field of cancer research
2007: Dickson Prize in Medicine
2007: Louisa Gross Horwitz Prize of Columbia University, shared with Elizabeth Blackburn and Joseph G. Gall
2008: Katharine Berkan Judd Award for Cancer Research
2008: The Pearl Meister Greengard Prize with Elizabeth Blackburn and Vicki Lundbald “for their insight into cellular ageing and cancer”10
2009: Nobel Prize in Physiology or Medicine, shared with Elizabeth Blackburn and Jack Szostak
2009: Paul Ehrlich and Ludwig Darmstaedter Prize, shared with Elizabeth Blackburn, “for their achievements in the discovery of telomeres and telomerase”
2010: Member of the Institute of Medicine
2013: Fellow of the American Association for Cancer Research (AACR)Academy
2016: Member of the American Philosophical Society



Journey Into Dyslexia (2011), dir. Susan Raymond, HBO, film. Documentary.



with E. H. Blackburn (1985), “Identification of a specific telomere terminal transferase activity in tetrahymena extracts”, Cell, 43, 405-13.

with E. H. Blackburn (1989),  “A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis”, Nature, 337, 331-37.

with Counter, C. M. / A. A. Avilion / C. E. LeFeuvre / N. G. Stewart. / C.B. Harley / S. Bacchetti (1992), “Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity”, EMBO J. , 11, 1921-29.

with Blasco, M. A. / H. W. Lee / P. M. Hande / E. Samper / P. M. Lansdorp / R. A. DePinho (1997), “Telomere Shortening and Tumor Formation by Mouse Cells Lacking Telomerase RNA”, Cell, 91, 25-34.

withKass-Eisler, A. (2000), “Recombination and telomeres”, TrendsBiochem. Sci, 25(4), 200-04.

with Chen, J.-L. / M. Blasco (2000), “A Secondary structure of vertebrate telomerase RNA”, Cell, 100, 503-14.

withHemann, M. T.  (2000), “Wild derived inbred mouse strains have short telomeres”,  Nucleic Acids Res., 28, 474-78.

with Chen, Q. / A. Ijpma, (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.

with W. Hemann, M. T. / L. Rudolph / M. Strong / R. A. DePinho / L. Chin  (2001), “Telomere dysfunction triggers developmentally regulated germ cell apoptosis”, Mol. Biol. Cell, 12, 2023-30.

with Hackett, J. (2001), “Telomere dysfunction increases mutation rate and genomic instability”, Cell, 106, 275-86.

with M. T., Strong / L. Y. Hao (2001), “The shortest telomere, not average telomere length, is critical for cell viability and chromosome stability”, Cell, 107, 67-77.

with IJpma, A. (2002),  “Short telomeres induce a DNA damage response in S. cerevisiae”, Mol. Biol. Cell, 14, 987-1001.

with Hackett, J.A. (2002), “Balancing instability: Dual roles for telomerase and telomere dysfunction in tumorigenesis”, Oncogene, 21, 619-26.

with Vidal-Cardenas, S.L. (2010), “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, 38(1), 60-71.

with Tom, H.I.(2010), “A sequence-dependent exonuclease activity from Tetrahymenathermophila”, BMC Biochem, 11, 45.

with Strong, M.A. / S.L. Vidal-Cardenas / B. Karim / H. Yu / N. Guo (2010), “Phenotypes in mTERT+/- and mTERT-/- mice are due to short telomeres, not telomere-independent functions of TERT ”, Mol. Cell Biol, 31, 2369-79.

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



“Carol W. Greider ”, John Hopkins Medicine, <>(last accessed 7 Jun. 2016).

Regents of the University of California (2009), ”‘Conversations with History: Carol W. Greider’ from Institute of International Studies”,  Berkeley university, <> (last accessed 13 Jun. 2016).

“Curriculum Vitae of Carol W. Greider”, Johns Hopkins University School of Medicine, <> (last accessed 13 Jun. 2016).

“Carol W. Greider – Biographical” (2015),, Nobel Media AB, <> (last accessed 13 Jun. 2016).

“Member’s profile (archive) Carol W. Greider” (2009), American Society for Cell Biology, <> (last accessed 13 Jun. 2016).

Nuzzo, Regina (2005), “Biography of Carol W. Greider”, The National Academy of Sciences, 102(23), 8077-79, <> (last accessed 13 Jun. 2016).

“Rosenstiel Award-Past Winners”, Brandeis University, <> (last accessed 13 Jun. 2016).



  1. “The Nobel Prize in Physiology or Medicine 2009 – Facts” (2014), Nobel Media AB ,<> (last accessed 9 Jan. 2017).
  2. (2015) “Carol W. Greider – Biographical ”(2014),, Nobel Media AB, : <> (last accessed 13 Jun. 2016).
  3. “Nathaniel Charles Comfort, M.S., Ph.D.”, Johns Hopkins Medicine, The Johns Hopkins University, The Johns Hopkins Hospital, and Johns Hopkins Health System, <> (last accessed 23 Feb. 2017).
  4. “Carol Greider, Ph.D.”, Academy of Achievement, American Academy of Achievement, <> (last accessed 23 Feb. 2017).
  5. “Member’s profile (archive) Carol W. Greider” (2009), American Society for Cell Biology, <> (last accessed 13 Jun. 2016).
  6. “Recipients of the Pearl Meister Greengard Prize” (2014), The Rockefeller University,<> (last accessed 14 Jun. 2016).
  7. “Rosential Award Past Winners” (2015), Brandeis University, <> (last accessed 13 Jun. 2016).
  8. “THE PASSANO AWARDS 1945-2015” (2015), The Passano Foundation, <> (last accessed 23 Feb. 2017).
  9. “Richard Lounsbery Award”, National Academy of Sciences,  <> (last accessed 13 Jun. 2016).
  10. “Recipients of the Pearl Meister Greengard Prize”, The Rockefeller University,<> (last accessed 14 Jun. 2016).
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