Carol W. Greider

{{Short description|American molecular biologist and Nobel laureate}}

{{Infobox scientist

| name = Carol W. Greider

| image = Carol Greider by Chris Michel 1s946948-11-23.jpg

| caption = Greider in 2021

| birth_name = Carolyn Widney Greider

| birth_date = {{Birth date and age|1961|04|15}}

| birth_place = San Diego, California, U.S.

| fields = Molecular biology

| workplaces = Cold Spring Harbor Laboratory
Johns Hopkins School of Medicine
University of California, Santa Cruz

| education = University of California, Santa Barbara (BA)
University of Göttingen
University of California, Berkeley (PhD)

| thesis_title = Identification of a specific telomere terminal transferase activity in Tetrahymena extracts

| thesis_url = https://pubmed.ncbi.nlm.nih.gov/3907856/

| thesis_year = 1985

| doctoral_advisor = Elizabeth Blackburn

| academic_advisors = Beatrice M. Sweeney
David J. Asai
Leslie Wilson

| known_for = Discovery of telomerase

| awards = Richard Lounsbery Award (2003)
Lasker Award (2006)
Louisa Gross Horwitz Prize (2007)
Nobel Prize in Physiology or Medicine (2009)

| spouse = {{Marriage|Nathaniel C. Comfort|1993|2011|end=div.}}

| children = 2

}}

Carolyn Widney Greider (born April 15, 1961) is an American molecular biologist and Nobel laureate. She is a Distinguished Professor of Molecular, Cell, and Developmental Biology{{Cite web|title=Eminent biologist Carol Greider to join UC Santa Cruz faculty|url=https://news.ucsc.edu/2020/05/carol-greider.html|last=Stephens|first=Tim|website=UC Santa Cruz News|language=en|access-date=May 22, 2020}} at the University of California, Santa Cruz.

Greider discovered the enzyme telomerase in 1984, while she was a graduate student of Elizabeth Blackburn at the University of California, Berkeley. Greider pioneered research on the structure of telomeres, the ends of the chromosomes. 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.{{Cite web |url=http://blogs.dnalc.org/dnaftb/2009/10/05/blackburn-greider-and-szostak-share-nobel-for-telomeres/ |title=Blackburn, Greider, and Szostak share Nobel |publisher=Dolan DNA Learning Center |access-date=October 5, 2009 |url-status=dead |archive-url=https://web.archive.org/web/20091022193507/http://blogs.dnalc.org/dnaftb/2009/10/05/blackburn-greider-and-szostak-share-nobel-for-telomeres/ |archive-date=October 22, 2009 }}

Early life and education

Greider was born in San Diego, California.[http://www.hopkinsmedicine.org/Press_releases/2008/10_01a_08.html Hopkins "Telomere" expert Carol Greider shares Germany's largest science prize] Her father, Kenneth Greider, was a physics professor.{{Cite web| title = Former Davis resident receives Nobel Prize| work = The California Aggie| access-date = April 7, 2015| date = October 12, 2009| url = http://www.theaggie.org/2009/10/12/former-davis-resident-receives-nobel-prize/}} Her family moved from San Diego to Davis, California, where she spent many of her early years and graduated from Davis Senior High School in 1979. She graduated from the College of Creative Studies at the University of California, Santa Barbara, with a B.A. in biology in 1983. During this time she also studied at the University of Göttingen and made significant discoveries there.[http://www.uni-goettingen.de/de/3240.html?cid=3435 Press release], University of Göttingen (December 9, 2009). (German)

Greider is dyslexic and states that her "compensatory skills also played a role in my success as a scientist because one has to intuit many different things that are going on at the same time and apply those to a particular problem".{{cite news|url=http://dyslexia.yale.edu/greider.html|title=Carol Greider, Scientist, Nobel Prize Winner |publisher=The Yale Center for Dyslexia & Creativity|last=Kathy Crockett|work=Yale University|access-date=March 5, 2015}} Greider initially suspected her dyslexia after seeing patterns of common mistakes such as backward words when she received back graded work in the first grade.{{Cite web|url=https://www.nobelprize.org/nobel_prizes/medicine/laureates/2009/greider-bio.html|title=Carol W. Greider – Biographical|website=www.nobelprize.org|access-date=September 28, 2017}} Greider started to memorize words and their spellings rather than attempting to sound out the spelling of words. Greider has worked significantly to overcome her dyslexia to become successful in her professional life and credits her dyslexia as helping her appreciate differences and making unusual decisions such as the one to work with Tetrahymena, an unusual organism.

Greider initially had difficulty getting into graduate school because of her low GRE scores due to her dyslexia. Greider applied to thirteen grad schools and was accepted to only two, California Institute of Technology and the University of California, Berkeley. She chose to study at Berkeley.

Discovery of telomerase

Greider completed her Ph.D. in molecular biology in 1987 at Berkeley under Elizabeth Blackburn. While at Berkeley, Greider and Blackburn discovered how chromosomes are protected by telomeres and the enzyme telomerase.{{Cite web| title = The Nobel Prize in Physiology or Medicine 2009| access-date = April 7, 2015| url = https://www.nobelprize.org/prizes/medicine/2009/illustrated-information/}}

Greider joined Blackburn's laboratory in April 1984 looking for the enzyme that was hypothesized to add extra DNA bases to the ends of chromosomes. Without the extra bases, which are added as repeats of a six-base pair motif, chromosomes are shortened during DNA replication, eventually resulting in chromosome deterioration and senescence or cancer-causing chromosome fusion. Blackburn and Greider looked for the enzyme in the model organism Tetrahymena thermophila, a fresh-water protozoan with a large number of telomeres.{{Cite journal| doi = 10.1073/pnas.0503019102| pmid = 15928079| year = 2005| last1 = Nuzzo | first1 = R.|author-link= Regina Nuzzo | title = Biography of Carol W. Greider| volume = 102| issue = 23| pages = 8077–8079| pmc = 1149435| journal = Proceedings of the National Academy of Sciences of the United States of America|bibcode = 2005PNAS..102.8077N | doi-access = free}}

On December 25, 1984, Greider obtained results indicating that a particular enzyme was likely responsible. After six months of additional research, Greider and Blackburn concluded that it was the enzyme responsible for telomere addition. They published their findings in the journal Cell in December 1985.{{Cite journal

| pmid = 3907856

| year = 1985

| last1 = Greider | first1 = C. W.

| last2 = Blackburn | first2 = E. H.

| title = Identification of a specific telomere terminal transferase activity in Tetrahymena extracts

| volume = 43

| issue = 2 Pt 1

| pages = 405–413

| journal = Cell

| doi = 10.1016/0092-8674(85)90170-9

| doi-access = free

}} The enzyme, originally called "telomere terminal transferase," is now known as telomerase. Telomerase rebuilds the tips of chromosomes and determines the life span of cells.{{Cite web| title = Carol Greider, Ph.D.| work = Johns Hopkins Medicine – Research – Awards – Nobel| access-date = April 7, 2015| url = http://www.hopkinsmedicine.org/research/awards/nobel/nobel_prize_greider.html| archive-date = August 28, 2015| archive-url = https://web.archive.org/web/20150828214812/http://www.hopkinsmedicine.org/research/awards/nobel/nobel_prize_greider.html| url-status = dead}}

Greider's additional research to confirm her discovery was largely focused on identifying the mechanism that telomerase uses for elongation.{{Cite news|last=Aicher |first=Toby |date=March 18, 2015|title=Science Spotlight: Nobel Laureate Carol Greider|work=The Middlebury Campus|url=https://middleburycampus.com/30712/arts-academics/science-spotlight-nobel-laureate-carol-greider/ |access-date=January 24, 2020}} Greider chose to use RNA degrading enzymes and saw that the telomeres stopped extending, which was an indication that RNA was involved in the enzyme.

Subsequent career

Greider then started her laboratory as a Cold Spring Harbor Laboratory Fellow, and also held a faculty position, at the Cold Spring Harbor Laboratory, Long Island, New York. Greider continued to study Tetrahymena telomerase, cloning the gene encoding the RNA component and demonstrating that it provided the template for the TTGGGG telomere repeats (1989){{cite journal |last1=Greider |first1=Carol W. |last2=Blackburn |first2=Elizabeth H. |s2cid=29191852 |title=A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis |journal=Nature |date=January 1989 |volume=337 |issue=6205 |pages=331–337 |doi=10.1038/337331a0|pmid=2463488 |bibcode=1989Natur.337..331G }} as well as establishing that telomerase is processive (1991).{{cite journal |last1=Greider |first1=C W |title=Telomerase is processive. |journal=Molecular and Cellular Biology |date=September 1991 |volume=11 |issue=9 |pages=4572–4580 |doi=10.1128/MCB.11.9.4572|pmid=1875940 |pmc=361337 }} She was also able to reconstitute Tetrahymena telomerase in vitro (1994){{cite journal |last1=Autexier |first1=C |last2=Greider |first2=C W |title=Functional reconstitution of wild-type and mutant Tetrahymena telomerase. |journal=Genes & Development |date=March 1, 1994 |volume=8 |issue=5 |pages=563–575 |doi=10.1101/gad.8.5.563|pmid=7523243 |doi-access=free }} and define the mechanisms of template utilization (1995).{{cite journal |last1=Autexier |first1=C |last2=Greider |first2=C W |title=Boundary elements of the Tetrahymena telomerase RNA template and alignment domains. |journal=Genes & Development |date=September 15, 1995 |volume=9 |issue=18 |pages=2227–2239 |doi=10.1101/gad.9.18.2227|pmid=7557377 |doi-access=free }} Greider also worked with Calvin Harley to show that telomere shortening underlies cellular senescence (1990).{{cite journal |last1=Greider |first1=Carol W. |title=Telomeres, telomerase and senescence |journal=BioEssays |date=August 1990 |volume=12 |issue=8 |pages=363–369 |doi=10.1002/bies.950120803|pmid=2241933 |s2cid=11920124 |doi-access=free }}{{cite journal |last1=Harley |first1=Calvin B. |last2=Futcher |first2=A. Bruce |last3=Greider |first3=Carol W. |s2cid=1145492 |title=Telomeres shorten during ageing of human fibroblasts |journal=Nature |date=May 1990 |volume=345 |issue=6274 |pages=458–460 |doi=10.1038/345458a0|pmid=2342578 |bibcode=1990Natur.345..458H }} To further test this idea mouse and human telomerase were characterized (1993){{cite journal |last1=Prowse |first1=K. R. |last2=Avilion |first2=A. A. |last3=Greider |first3=C. W. |title=Identification of a nonprocessive telomerase activity from mouse cells. |journal=Proceedings of the National Academy of Sciences |date=February 15, 1993 |volume=90 |issue=4 |pages=1493–1497 |doi=10.1073/pnas.90.4.1493 |pmid=8434010 |pmc=45900 |bibcode=1993PNAS...90.1493P |doi-access=free }} (1995){{cite journal |last1=Feng |first1=J. |last2=Funk |first2=W. |last3=Wang |first3=S. |last4=Weinrich |first4=S. |last5=Avilion |first5=A. |last6=Chiu |first6=C. |last7=Adams |first7=R. |last8=Chang |first8=E. |last9=Allsopp |first9=R. |last10=Yu |first10=J. |last11=al. |first11=e. |title=The RNA component of human telomerase |journal=Science |date=September 1, 1995 |volume=269 |issue=5228 |pages=1236–1241 |doi=10.1126/science.7544491|pmid=7544491 |bibcode=1995Sci...269.1236F |s2cid=9440710 }} and the mouse telomerase RNA component was cloned (1995).{{cite journal |last1=Blasco |first1=M. |last2=Funk |first2=W. |last3=Villeponteau |first3=B. |last4=Greider |first4=C. |title=Functional characterization and developmental regulation of mouse telomerase RNA |journal=Science |date=September 1, 1995 |volume=269 |issue=5228 |pages=1267–1270 |doi=10.1126/science.7544492|pmid=7544492 |bibcode=1995Sci...269.1267B |s2cid=1315745 }}

During this time, Greider, in collaboration with Ronald A. DePinho, produced the first telomerase knockout mouse,{{cite journal |last1=Blasco |first1=María A |last2=Lee |first2=Han-Woong |last3=Hande |first3=M.Prakash |last4=Samper |first4=Enrique |last5=Lansdorp |first5=Peter M |last6=DePinho |first6=Ronald A |last7=Greider |first7=Carol W |title=Telomere Shortening and Tumor Formation by Mouse Cells Lacking Telomerase RNA |journal=Cell |date=October 1997 |volume=91 |issue=1 |pages=25–34 |doi=10.1016/s0092-8674(01)80006-4|pmid=9335332 |s2cid=13366934 |doi-access=free }} showing that although telomerase is dispensable for life, increasingly short telomeres result in various deleterious phenotypes, colloquially referred to as premature aging.{{cite journal |last1=Rudolph |first1=Karl Lenhard |last2=Chang |first2=Sandy |last3=Lee |first3=Han-Woong |last4=Blasco |first4=Maria |last5=Gottlieb |first5=Geoffrey J |last6=Greider |first6=Carol |last7=DePinho |first7=Ronald A |title=Longevity, Stress Response, and Cancer in Aging Telomerase-Deficient Mice |journal=Cell |date=March 1999 |volume=96 |issue=5 |pages=701–712 |doi=10.1016/s0092-8674(00)80580-2|pmid=10089885 |s2cid=11991355 |doi-access=free }} In the mid-1990s, Greider was recruited by Michael D. West, founder of biotechnology company Geron (now CEO of AgeX Therapeutics) to join the company's Scientific Advisory Board{{cite web|title=Geron Corporation 10K 1996|url= http://www.bio.net/bionet/mm/ageing/1996-May/002331.html}} and remained on the Board until 1997.

Greider accepted a faculty position at the Johns Hopkins University School of Medicine in 1997. Greider continued to study telomerase deficient mice and saw that her sixth generation of mice had become entirely sterile,{{cite journal |last1=Lee |first1=Han-Woong |last2=Blasco |first2=Maria A. |last3=Gottlieb |first3=Geoffrey J. |last4=Horner |first4=James W. |last5=Greider |first5=Carol W. |last6=DePinho |first6=Ronald A. |s2cid=4385788 |title=Essential role of mouse telomerase in highly proliferative organs |journal=Nature |date=April 1998 |volume=392 |issue=6676 |pages=569–574 |doi=10.1038/33345|pmid=9560153 |bibcode=1998Natur.392..569L }} but when mated with control mice the telomerase deficient mice were able to regenerate their telomeres.{{cite journal |last1=Hemann |first1=Michael T |last2=Strong |first2=Margaret A |last3=Hao |first3=Ling-Yang |last4=Greider |first4=Carol W |title=The Shortest Telomere, Not Average Telomere Length, Is Critical for Cell Viability and Chromosome Stability |journal=Cell |date=October 2001 |volume=107 |issue=1 |pages=67–77 |doi=10.1016/s0092-8674(01)00504-9|pmid=11595186 |s2cid=10719526 |doi-access=free }} Greider continued to work on telomerase biochemistry, defining the secondary structure (2000) {{cite journal |last1=Chen |first1=Jiunn-Liang |last2=Blasco |first2=Maria A |last3=Greider |first3=Carol W |title=Secondary Structure of Vertebrate Telomerase RNA |journal=Cell |date=March 2000 |volume=100 |issue=5 |pages=503–514 |doi=10.1016/s0092-8674(00)80687-x|pmid=10721988 |s2cid=15642776 |doi-access=free }} and template boundary (2003){{cite journal |last1=Chen |first1=J.-L. |title=Template boundary definition in mammalian telomerase |journal=Genes & Development |date=November 15, 2003 |volume=17 |issue=22 |pages=2747–2752 |doi=10.1101/gad.1140303|pmid=14630939 |pmc=280623 }} of vertebrate telomerase RNA as well as analyzing the pseudoknot structure in human telomerase RNA (2005).{{cite journal |last1=Chen |first1=J.-L. |last2=Greider |first2=C. W. |title=Functional analysis of the pseudoknot structure in human telomerase RNA |journal=Proceedings of the National Academy of Sciences |date=April 22, 2005 |volume=102 |issue=23 |pages=8080–8085 |doi=10.1073/pnas.0502259102|pmid=15849264 |pmc=1149427 |bibcode=2005PNAS..102.8080C |doi-access=free }} In addition to working in Tetrahymena and mammalian systems, Greider also studied telomeres and telomerase in the yeast Saccharomyces cerevisiae, further characterizing the recombination-based gene conversion mechanism that yeast cells null for telomerase use to maintain telomeres (1999){{cite journal |last1=Le |first1=S |last2=Moore |first2=JK |last3=Haber |first3=JE |last4=Greider |first4=CW |title=RAD50 and RAD51 define two pathways that collaborate to maintain telomeres in the absence of telomerase. |journal=Genetics |date=May 1999 |volume=152 |issue=1 |pages=143–52 |doi=10.1093/genetics/152.1.143 |pmid=10224249|pmc=1460580 }} (2001).{{cite journal |last1=Chen |first1=Q. |last2=Ijpma |first2=A. |last3=Greider |first3=C. W. |title=Two Survivor Pathways That Allow Growth in the Absence of Telomerase Are Generated by Distinct Telomere Recombination Events |journal=Molecular and Cellular Biology |date=March 1, 2001 |volume=21 |issue=5 |pages=1819–1827 |doi=10.1128/MCB.21.5.1819-1827.2001|pmid=11238918 |pmc=86745 }} Greider also showed that short telomeres elicit a DNA damage response in yeast (2003).{{cite journal |last1=IJpma |first1=Arne S. |last2=Greider |first2=Carol W. |last3=Koshland |first3=Douglas |title=Short Telomeres Induce a DNA Damage Response in |journal=Molecular Biology of the Cell |date=March 2003 |volume=14 |issue=3 |pages=987–1001 |doi=10.1091/mbc.02-04-0057|pmid=12631718 |pmc=151574 }}

Greider, Blackburn, and Szostak shared the 2006 Albert Lasker Award for Basic Medical Research for their work on telomeres,{{cite web|title="Telomere" Expert Carol Greider Shares 2009 Nobel Prize in Physiology or Medicine|url=http://www.hopkinsmedicine.org/news/media/releases/telomere_expert_carol_greider_shares_2009_nobel_prize_in_physiology_or_medicine|website=Johns Hopkins University|access-date=March 13, 2015}} before jointly receiving the Nobel Prize in 2009.

In February 2014, Greider was named a Bloomberg Distinguished Professor at Johns Hopkins University.{{cite news |url=http://hub.jhu.edu/2014/02/17/bloomberg-distinguished-professors |title=With Bloomberg Distinguished Professorships, Johns Hopkins aims to foster cross-specialty collaboration |last=Brooks |first=Kelly |date=February 17, 2014 |website = Hub |publisher=Johns Hopkins University |access-date=March 12, 2015}}

Greider served as director of and professor at the Department of Molecular Biology and Genetics at Johns Hopkins Medicine. Greider was first promoted to Daniel Nathans Professor at the Department of Molecular Biology and Genetics in 2004.{{Cite web|url=http://www.womenofhopkins.com/greider|title=The Women of Hopkins|website=The Women of Hopkins|access-date=October 5, 2017}}

As of 2021, she is Distinguished Professor of Molecular, Cellular, and Developmental Biology at UC Santa Cruz.{{Cite web |last=Stephens |first=Tim |title=Eminent biologist Carol Greider to join UC Santa Cruz faculty |url=https://news.ucsc.edu/2020/05/carol-greider.html |access-date=2025-01-04 |website=UC Santa Cruz News |language=en}}

Greider's lab employs both student and post-doctoral trainees{{Cite news|url=https://www.wsj.com/articles/SB10001424127887323978104578335013555368892|title=NIH Cuts Began Ahead of Sequester|last=Paletta|first=Thomas M. Burton and Damian|date=March 2, 2013|work=Wall Street Journal|access-date=October 5, 2017|language=en-US|issn=0099-9660}} to further examine the relationships between the biology of telomeres and their connection to disease. Greider's lab uses a variety of tools including yeast, mice, and biochemistry to look at progressive telomere shortening.{{Cite web|url=http://www.greiderlab.org/index.html|title=The Greider Lab {{!}} Johns Hopkins Medicine|website=www.greiderlab.org|access-date=October 6, 2017}} Greider's lab is also researching how tumor reformation can be controlled by the presence of short telomeres. The lab's future work will focus more on identifying the processing and regulation of telomeres and telomere elongation.

Personal life

Greider married Nathaniel C. Comfort, a fellow academic, in 1992. They divorced in 2011. She has two children.{{cite news|url=http://artsandsciences.colorado.edu/magazine/2010/03/having-it-all-plus-doing-it-all/|title='Having it all' plus 'doing it all'|last=Clint Talbott|work=Colorado Arts & Sciences Magazine|access-date=March 5, 2015|archive-url=https://web.archive.org/web/20150220192119/http://artsandsciences.colorado.edu/magazine/2010/03/having-it-all-plus-doing-it-all/|archive-date=February 20, 2015|url-status=dead}}

Awards and honors

Pew Scholar in the Biomedical Sciences (1990–1994){{cite web |title=Carol W. Greider, Ph.D. |url=https://www.pewtrusts.org/en/projects/pew-biomedical-scholars/directory-of-pew-scholars/1990/carol-greider |website=bit.ly |access-date=January 31, 2020}}
Gairdner Foundation International Award (1998){{cite web |title=Carol Greider |url=https://gairdner.org/award_winners/carol-greider/ |website=Gairdner Foundation |access-date=January 31, 2020}}
Rosenstiel Award in Basic Medical Research (1998) (shared with Elizabeth Blackburn){{cite web |title=Past Winners |url=https://www.brandeis.edu/rosenstiel/rosenstiel-award/past.html |website=www.brandeis.edu |access-date=January 31, 2020 |language=en}}
Member of the American Society for Cell Biology (1999){{citation needed|date=April 2021}}
Passano Foundation Award (1999) (shared with Elizabeth Blackburn){{cite web |title=Recipients of the Passano Laureate and Physician Scientist Awards |url=https://www.passanofoundation.org/services |website=The Passano Foundation, Inc. |access-date=January 31, 2020}}
Harvey Society Lecture (2000){{citation needed|date=April 2021}}
Academy of Achievement Golden Plate Award (2000){{cite web|title=Carol W. Greider Biography and Interview |website=www.achievement.org|publisher=American Academy of Achievement|date=June 16, 2000|url=https://www.achievement.org/achiever/carol-greider-ph-d/#interview}}
Fellow of the American Academy of Arts and Sciences (2003){{cite web|title=Book of Members, 1780–2010: Chapter B|url=http://www.amacad.org/publications/BookofMembers/ChapterB.pdf|publisher=American Academy of Arts and Sciences|access-date=June 9, 2011}}
Member of the National Academy of Sciences (2003){{cite web|title=Greider, Carol W.|url=http://www.nasonline.org/site/Dir/705195530?pg=vprof&mbr=1003047|work=National Academy of Sciences|access-date=June 9, 2011}}
Richard Lounsbery Award (2003), National Academy of Sciences[http://www.nasonline.org/site/PageServer?pagename=AWARDS_lounsbery NAS Online] {{Webarchive|url=https://web.archive.org/web/20061209124335/http://www.nasonline.org/site/PageServer?pagename=AWARDS_lounsbery |date=December 9, 2006 }} ("For her pioneering biochemical and genetic studies of telomerase, the enzyme that maintains the ends of chromosomes in eukaryotic cells.")
Member of the American Society for Biochemistry and Molecular Biology (2004){{citation needed|date=April 2021}}
Lila Gruber Cancer Research Award (2006){{cite web |title=Lila and Murray Gruber Memorial Cancer Research Award and Lectureship |url=https://www.aad.org/member/career/awards/gruber |website=www.aad.org |access-date=January 31, 2020}}
Albert Lasker Award for Basic Medical Research (2006) (shared with Elizabeth Blackburn and Jack Szostak){{cite web |last1=Foundation |first1=Lasker |title=2006 Lasker Awards |url=http://www.laskerfoundation.org/awards/year/2006/ |website=The Lasker Foundation |access-date=January 31, 2020 |language=en}}
Wiley Prize in Biomedical Sciences (2006) (shared with Elizabeth Blackburn){{cite web |title=Wiley: The Wiley Foundation Announces Recipients of the Fifth Annual Wiley Prize in Biomedical Sciences |url=https://www.wiley.com/WileyCDA/PressRelease/pressReleaseId-49048.html |website=www.wiley.com |access-date=January 31, 2020}}
Dickson Prize in Medicine (2007){{cite web |title=Carol W. Greider, PhD {{!}} Dickson Prize in Medicine {{!}} University of Pittsburgh |url=http://www.dicksonprize.pitt.edu/recipients/2007-greider.php |website=www.dicksonprize.pitt.edu |access-date=January 31, 2020}}
Louisa Gross Horwitz Prize of Columbia University (2007) (shared with Elizabeth Blackburn and Joseph G. Gall){{cite web |title=Horwitz Prize Awardees |url=https://www.cuimc.columbia.edu/research/louisa-gross-horwitz-prize/horwitz-prize-awardees |website=Columbia University Irving Medical Center |access-date=January 31, 2020 |language=en |date=June 20, 2018}}
The Pearl Meister Greengard Prize (2008) (shared with Elizabeth Blackburn and Vicki Lundblad){{cite web |title=Recipients |url=https://www.rockefeller.edu/greengard-prize/recipients/ |website=Greengard Prize |access-date=January 31, 2020 |language=en}}
Nobel Prize in Physiology or Medicine (2009) (shared with Elizabeth Blackburn and Jack Szostak)"{{Cite web|url=http://nobelprize.org/nobel_prizes/medicine/laureates/2009/|title=Nobel Prize in Physiology or Medicine 2009|publisher=Nobel Foundation|access-date=October 5, 2009}}
Paul Ehrlich and Ludwig Darmstaedter Prize (2009) (shared with Elizabeth Blackburn){{cite web |title=Press Release 2008 |url=https://www.uni-frankfurt.de/44563345/PressemitPaulEhrlichPreis2009-engl_.pdf |access-date=January 31, 2020}}
Member of the Institute of Medicine (2010){{cite web|title=IOM Class of 2010|url=http://www.iom.edu/About-IOM/Membership/IOMClassof2010.aspx|work=Institute of Medicine|access-date=June 9, 2011|archive-url=https://web.archive.org/web/20110422005017/http://iom.edu/About-IOM/Membership/IOMClassof2010.aspx|archive-date=April 22, 2011|url-status=dead}}{{cite news|title=Carson, Hopkins Colleagues Named to Institute of Medicine|url=http://www.hopkinschildrens.org/Carson-Hopkins-Colleagues-Named-to-Institute-of-Medicine.aspx|publisher=hopkinschildrens.org|date=October 11, 2010|access-date=October 1, 2013|archive-date=November 25, 2010|archive-url=https://web.archive.org/web/20101125032422/http://www.hopkinschildrens.org/Carson-Hopkins-Colleagues-Named-to-Institute-of-Medicine.aspx|url-status=dead}}
Member, American Philosophical Society (2016){{cite web |title=APS Member History |url=https://search.amphilsoc.org/memhist/search?creator=Greider&title=&subject=&subdiv=&mem=&year=&year-max=&dead=&keyword=&smode=advanced |website=search.amphilsoc.org |access-date=January 31, 2020}}
Pinnacle Award (2019), Association for Women in Science{{cite web |title=Press Release AWIS 2019 Awards |url=https://www.awis.org/innovation-inclusion-summit-awards-carol-greider-mareena-robinson-snowden/ |website=www.awis.org|date=November 8, 2018 }}
Award for Excellence in Molecular Diagnostics, Association for Molecular Pathology (2022){{Cite web |title=Past Recipients |url=https://www.amp.org/membership/awards-grants-honors/amp-award-for-excellence-in-molecular-diagnostics/past-recipients/ |access-date=2023-04-12 |website=Association for Molecular Pathology |language=en}}

Selected works

{{Cite journal|doi=10.1016/0092-8674(85)90170-9 |last1=Greider |first1=C. W. |last2=Blackburn |first2=E. H. |name-list-style=amp |year=1985 |title=Identification of a specific telomere terminal transferase activity in Tetrahymena extracts |journal=Cell |volume=43 |issue=2 Pt. 1 |pages=405–413 |pmid=3907856 |doi-access=free }}
{{Cite journal|last1=Greider |first1=C. W. |last2=Blackburn |first2=E. H. |name-list-style=amp |year=1996 |title=Telomeres, Telomerase and Cancer |journal=Scientific American |volume= 274|issue= 2|pages=92–97 |doi=10.1038/scientificamerican0296-92 |pmid=8560215 |bibcode=1996SciAm.274b..92G }}