Carolyn Widney Greider (born April 15, 1961) is an American
molecular biologist
Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
and Nobel laureate. She is a Distinguished Professor of
Molecular
A molecule is a group of two or more atoms that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...
,
Cell, and
Developmental Biology
Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of Regeneration (biology), regeneration, asexual reproduction, metamorphosis, and the growth and di ...
at the
University of California, Santa Cruz
The University of California, Santa Cruz (UC Santa Cruz or UCSC) is a public university, public Land-grant university, land-grant research university in Santa Cruz, California, United States. It is one of the ten campuses in the University of C ...
.
Greider discovered the
enzyme
An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
telomerase
Telomerase, also called terminal transferase, is a ribonucleoprotein that adds a species-dependent telomere repeat sequence to the 3' end of telomeres. A telomere is a region of repetitive sequences at each end of the chromosomes of most euka ...
in 1984, while she was a graduate student of
Elizabeth Blackburn
Elizabeth Helen Blackburn (born 26 November 1948) is an Australian-American Nobel laureate who is the former president of the Salk Institute for Biological Studies. In 1984, Blackburn co-discovered telomerase, the enzyme that replenishes the ...
at the
University of California, Berkeley
The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
. Greider pioneered research on the structure of
telomere
A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes (see #Sequences, Sequences). Telomeres are a widespread genetic feature most commonly found in eukaryotes. In ...
s, the ends of the
chromosome
A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s. She was awarded the 2009
Nobel Prize for Physiology or Medicine
The Nobel Prize in Physiology or Medicine () is awarded yearly by the Nobel Assembly at the Karolinska Institute, Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single ...
, along with Blackburn and
Jack W. Szostak, for their discovery that telomeres are protected from progressive shortening by the enzyme
telomerase
Telomerase, also called terminal transferase, is a ribonucleoprotein that adds a species-dependent telomere repeat sequence to the 3' end of telomeres. A telomere is a region of repetitive sequences at each end of the chromosomes of most euka ...
.
Early life and education
Greider was born in
San Diego
San Diego ( , ) is a city on the Pacific coast of Southern California, adjacent to the Mexico–United States border. With a population of over 1.4 million, it is the List of United States cities by population, eighth-most populous city in t ...
,
California
California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
. Her father, Kenneth Greider, was a
physics
Physics is the scientific study of matter, its Elementary particle, fundamental constituents, its motion and behavior through space and time, and the related entities of energy and force. "Physical science is that department of knowledge whi ...
professor. Her family moved from San Diego to
Davis, California
Davis is the most populous city in Yolo County, California, United States. Located in the Sacramento Valley region of Northern California, the city had a population of 66,850 in 2020, not including the on-campus population of the University of ...
, 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
The College of Creative Studies (CCS) is the smallest of the three Undergraduate education, undergraduate colleges at the University of California, Santa Barbara, unique within the University of California system in terms of structure and phi ...
at the
University of California, Santa Barbara
The University of California, Santa Barbara (UC Santa Barbara or UCSB) is a Public university, public Land-grant university, land-grant research university in Santa Barbara County, California, United States. Tracing its roots back to 1891 as an ...
, with a
B.A. in
biology
Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
in 1983. During this time she also studied at the
University of Göttingen
The University of Göttingen, officially the Georg August University of Göttingen (, commonly referred to as Georgia Augusta), is a Public university, public research university in the city of Göttingen, Lower Saxony, Germany. Founded in 1734 ...
and made significant discoveries there.
Greider is
dyslexic
Dyslexia (), previously known as word blindness, is a learning disability that affects either reading or writing. Different people are affected to different degrees. Problems may include difficulties in spelling words, reading quickly, writ ...
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".
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. 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
The California Institute of Technology (branded as Caltech) is a private research university in Pasadena, California, United States. The university is responsible for many modern scientific advancements and is among a small group of institutes ...
and the
University of California, Berkeley
The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
.
She chose to study at Berkeley.
Discovery of telomerase
Greider completed her Ph.D. in
molecular biology
Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
in 1987 at
Berkeley under
Elizabeth Blackburn
Elizabeth Helen Blackburn (born 26 November 1948) is an Australian-American Nobel laureate who is the former president of the Salk Institute for Biological Studies. In 1984, Blackburn co-discovered telomerase, the enzyme that replenishes the ...
. While at Berkeley, Greider and Blackburn discovered how chromosomes are protected by
telomere
A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes (see #Sequences, Sequences). Telomeres are a widespread genetic feature most commonly found in eukaryotes. In ...
s and the enzyme
telomerase
Telomerase, also called terminal transferase, is a ribonucleoprotein that adds a species-dependent telomere repeat sequence to the 3' end of telomeres. A telomere is a region of repetitive sequences at each end of the chromosomes of most euka ...
.
Greider joined Blackburn's laboratory in April 1984 looking for the enzyme that was hypothesized to add extra
DNA bases to the ends of
chromosome
A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s. Without the extra bases, which are added as repeats of a six-base pair motif, chromosomes are shortened during
DNA replication
In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all life, living organisms, acting as the most essential part of heredity, biolog ...
, eventually resulting in chromosome deterioration and
senescence
Senescence () or biological aging is the gradual deterioration of Function (biology), functional characteristics in living organisms. Whole organism senescence involves an increase in mortality rate, death rates or a decrease in fecundity with ...
or cancer-causing chromosome fusion. Blackburn and Greider looked for the enzyme in the model organism ''
Tetrahymena thermophila'', a fresh-water
protozoa
Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
n with a large number of telomeres.
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. 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.
Greider's additional research to confirm her discovery was largely focused on identifying the mechanism that telomerase uses for elongation.
Greider chose to use
RNA
Ribonucleic acid (RNA) is a polymeric molecule that is essential for most biological functions, either by performing the function itself (non-coding RNA) or by forming a template for the production of proteins (messenger RNA). RNA and deoxyrib ...
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
Cold Spring Harbor Laboratory (CSHL) is a private, non-profit institution with research programs focusing on cancer, neuroscience, botany, genomics, and quantitative biology. It is located in Laurel Hollow, New York, in Nassau County, on ...
,
Long Island, New York
Long Island is a densely populated continental island in southeastern New York (state), New York state, extending into the Atlantic Ocean. It constitutes a significant share of the New York metropolitan area in both population and land are ...
. 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) as well as establishing that telomerase is processive (1991). She was also able to reconstitute ''Tetrahymena'' telomerase ''in vitro'' (1994) and define the mechanisms of template utilization (1995). Greider also worked with Calvin Harley to show that telomere shortening underlies cellular senescence (1990). To further test this idea mouse and human telomerase were characterized (1993) (1995) and the mouse telomerase RNA component was cloned (1995).
During this time, Greider, in collaboration with
Ronald A. DePinho, produced the first telomerase
knockout mouse
A knockout mouse, or knock-out mouse, is a genetically modified mouse (''Mus musculus'') in which researchers have inactivated, or " knocked out", an existing gene by replacing it or disrupting it with an artificial piece of DNA. They are importan ...
, showing that although telomerase is dispensable for life, increasingly short telomeres result in various deleterious
phenotype
In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
s, colloquially referred to as premature aging. 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 and remained on the Board until 1997.
Greider accepted a faculty position at the
Johns Hopkins University School of Medicine
The Johns Hopkins University School of Medicine (JHUSOM) is the medical school of Johns Hopkins University, a Private university, private research university in Baltimore, Maryland. Established in 1893 following the construction of the Johns Ho ...
in 1997. Greider continued to study telomerase deficient mice and saw that her sixth generation of mice had become entirely sterile, but when mated with control mice the telomerase deficient mice were able to regenerate their
telomere
A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes (see #Sequences, Sequences). Telomeres are a widespread genetic feature most commonly found in eukaryotes. In ...
s.
Greider continued to work on telomerase biochemistry, defining the secondary structure (2000) and template boundary (2003) of vertebrate telomerase RNA as well as analyzing the pseudoknot structure in human telomerase RNA (2005). In addition to working in ''Tetrahymena'' and mammalian systems, Greider also studied telomeres and telomerase in the yeast ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungal microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have be ...
'', further characterizing the recombination-based gene conversion mechanism that yeast cells null for telomerase use to maintain telomeres (1999) (2001). Greider also showed that short telomeres elicit a DNA damage response in yeast (2003).
Greider, Blackburn, and Szostak shared the 2006
Albert Lasker Award for Basic Medical Research
The Albert Lasker Award for Basic Medical Research is one of the Lasker Award, prizes awarded by the Lasker Foundation for a fundamental discovery that opens up a new area of biomedical science. The award frequently precedes a Nobel Prize in Phys ...
for their work on telomeres, before jointly receiving the
Nobel Prize
The Nobel Prizes ( ; ; ) are awards administered by the Nobel Foundation and granted in accordance with the principle of "for the greatest benefit to humankind". The prizes were first awarded in 1901, marking the fifth anniversary of Alfred N ...
in 2009.
In February 2014, Greider was named a
Bloomberg Distinguished Professor at
Johns Hopkins University
The Johns Hopkins University (often abbreviated as Johns Hopkins, Hopkins, or JHU) is a private university, private research university in Baltimore, Maryland, United States. Founded in 1876 based on the European research institution model, J ...
.
Greider served as director of and professor at the Department of Molecular Biology and Genetics at
Johns Hopkins Medicine
The Johns Hopkins University School of Medicine (JHUSOM) is the medical school of Johns Hopkins University, a private research university in Baltimore, Maryland. Established in 1893 following the construction of the Johns Hopkins Hospital, the ...
.
Greider was first promoted to Daniel Nathans Professor at the Department of Molecular Biology and Genetics in 2004.
As of 2021, she is Distinguished Professor of Molecular, Cellular, and Developmental Biology at
UC Santa Cruz
The University of California, Santa Cruz (UC Santa Cruz or UCSC) is a public land-grant research university in Santa Cruz, California, United States. It is one of the ten campuses in the University of California system. Located in Monterey Bay ...
.
Greider's lab employs both student and post-doctoral trainees 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
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
,
mice
A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
, and biochemistry to look at progressive telomere shortening.
Greider's lab is also researching how
tumor
A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
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.
Awards and honors
*
Pew Scholar in the Biomedical Sciences (1990–1994)
*
Gairdner Foundation International Award (1998)
* Rosenstiel Award in Basic Medical Research (1998) (shared with Elizabeth Blackburn)
* Member of the
American Society for Cell Biology (1999)
* Passano Foundation Award (1999) (shared with Elizabeth Blackburn)
* Harvey Society Lecture (2000)
*
Academy of Achievement Golden Plate Award (2000)
* Fellow of the
American Academy of Arts and Sciences
The American Academy of Arts and Sciences (The Academy) is one of the oldest learned societies in the United States. It was founded in 1780 during the American Revolution by John Adams, John Hancock, James Bowdoin, Andrew Oliver, and other ...
(2003)
* Member of the
National Academy of Sciences
The National Academy of Sciences (NAS) is a United States nonprofit, NGO, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the ...
(2003)
*
Richard Lounsbery Award (2003), National Academy of Sciences
* Member of the
American Society for Biochemistry and Molecular Biology
The American Society for Biochemistry and Molecular Biology (ASBMB) is a learned society that was founded on December 26, 1906, at a meeting organized by John Jacob Abel (Johns Hopkins University). The roots of the society were in the American Ph ...
(2004)
* Lila Gruber Cancer Research Award (2006)
*
Albert Lasker Award for Basic Medical Research
The Albert Lasker Award for Basic Medical Research is one of the Lasker Award, prizes awarded by the Lasker Foundation for a fundamental discovery that opens up a new area of biomedical science. The award frequently precedes a Nobel Prize in Phys ...
(2006) (shared with
Elizabeth Blackburn
Elizabeth Helen Blackburn (born 26 November 1948) is an Australian-American Nobel laureate who is the former president of the Salk Institute for Biological Studies. In 1984, Blackburn co-discovered telomerase, the enzyme that replenishes the ...
and
Jack Szostak)
*
Wiley Prize in Biomedical Sciences (2006) (shared with Elizabeth Blackburn)
*
Dickson Prize in Medicine (2007)
*
Louisa Gross Horwitz Prize
The Louisa Gross Horwitz Prize for Biology or Biochemistry is an annual prize awarded by Columbia University to a researcher or group of researchers who have made an outstanding contribution in basic research in the fields of biology or biochemist ...
of
Columbia University
Columbia University in the City of New York, commonly referred to as Columbia University, is a Private university, private Ivy League research university in New York City. Established in 1754 as King's College on the grounds of Trinity Churc ...
(2007) (shared with Elizabeth Blackburn and
Joseph G. Gall)
* The Pearl Meister Greengard Prize (2008) (shared with Elizabeth Blackburn and Vicki Lundblad)
*
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine () is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, acco ...
(2009) (shared with Elizabeth Blackburn and Jack Szostak)"
*
Paul Ehrlich and Ludwig Darmstaedter Prize
The Paul Ehrlich and Ludwig Darmstaedter Prize is an annual award bestowed by the since 1952 for research in medicine. It carries a monetary prize of 120,000 Euro. The prize ceremony is traditionally held on the 14th of March, the birthday of Nob ...
(2009) (shared with Elizabeth Blackburn)
* Member of the
Institute of Medicine
The National Academy of Medicine (NAM), known as the Institute of Medicine (IoM) until 2015, is an American nonprofit, non-governmental organization. The National Academy of Medicine is a part of the National Academies of Sciences, Engineerin ...
(2010)
* Member,
American Philosophical Society
The American Philosophical Society (APS) is an American scholarly organization and learned society founded in 1743 in Philadelphia that promotes knowledge in the humanities and natural sciences through research, professional meetings, publicat ...
(2016)
* Pinnacle Award (2019),
Association for Women in Science
* Award for Excellence in Molecular Diagnostics,
Association for Molecular Pathology (2022)
Selected works
*
*
See also
*
List of female Nobel laureates
The Nobel Prizes are five separate prizes that, according to Alfred Nobel#Nobel Prize, Alfred Nobel's will of 1895, are awarded to "those who, during the preceding year, have conferred the greatest benefit to Mankind." Additionally, the Nobel Mem ...
*
List of RNA biologists
*
History of RNA biology
*
Timeline of women in science
This is a timeline of women in science, spanning from ancient history up to the 21st century. While the timeline primarily focuses on women involved with natural sciences such as astronomy, biology, chemistry and physics, it also includes women f ...
References
Further reading
*
External links
Greider Lab*
{{DEFAULTSORT:Greider, Carol
1961 births
American geneticists
American Nobel laureates
Fellows of the American Academy of Arts and Sciences
Johns Hopkins University faculty
Living people
Members of the National Academy of Medicine
Members of the United States National Academy of Sciences
Nobel laureates in Physiology or Medicine
Richard-Lounsbery Award laureates
People from Davis, California
People from San Diego
University of California, Berkeley alumni
University of California, Santa Barbara alumni
American women molecular biologists
Women Nobel laureates
Recipients of the Albert Lasker Award for Basic Medical Research
Davis Senior High School (California) alumni
American molecular biologists
20th-century American biologists
20th-century American women scientists
21st-century American biologists
21st-century American women scientists
Scientists with dyslexia
American scientists with disabilities
Fellows of the American Academy of Microbiology
Biologists from California
Members of the American Philosophical Society