View source for Barbara McClintock

{{Short description|American scientist and cytogeneticist (1902–1992)}}
{{About|the American scientist|the illustrator|Barbara McClintock (illustrator)}}
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{{Use American English|date=June 2023}}
{{Use mdy dates|date=September 2024}}
{{Infobox scientist
| name              = Barbara McClintock
| honorific_suffix  = 
| image             = Barbara McClintock (1902-1992) shown in her laboratory in 1947.jpg
| landscape         = yes
| caption           = McClintock in her laboratory, 1947
| birth_name        = Eleanor McClintock
| birth_date        = {{birth date|1902|6|16}}
| birth_place       = [[Hartford, Connecticut]], United States
| death_date        = {{death date and age|1992|9|2|1902|6|16}}
| death_place       = [[Huntington, New York]], United States
| field             = [[Cytogenetics]] (genetics scientist)
| work_institution  = {{plainlist|
* [[University of Missouri]]
* [[California Institute of Technology]]
* [[Cold Spring Harbor Laboratory]]
}}
| alma_mater        = [[Cornell University]] ([[B.S.|BS]], [[M.S.|MS]], PhD)
| doctoral_advisor  = 
| doctoral_students = 
| thesis_title      = A Cytological and Genetical Study of Triploid Maize
| thesis_year       = 1927
| thesis_url        = http://cornell.worldcat.org/title/cytological-and-genetical-study-of-triploid-maize/oclc/64683107
| known_for         = {{no wrap|[[Cytogenetics#McClintock's work on maize|Work in genetic structure of maize]]}}<br>[[Breakage-fusion-bridge cycle]]<br>[[Chromosomal crossover]]<br>[[DNA transposon]]<br>[[Mobile genetic elements]]<br>[[Transposase]]<br>[[Transposable element]]
| prizes            = {{Plainlist|
* [[Member of the National Academy of Sciences]] (1944)<ref>{{cite web|url=http://www.nasonline.org/member-directory/deceased-members/52803.html|title=Barbara McClintock|website=Nasonline.org|access-date=April 25, 2019}}</ref>
* [[National Medal of Science]] (1970)
* [[Thomas Hunt Morgan Medal]] (1981)
* [[Wolf Prize in Medicine]] (1981)
* [[Louisa Gross Horwitz Prize]] (1982)
* [[Nobel Prize in Physiology or Medicine]] (1983)}}
| footnotes         = 
| signature         = Barbara McClintock (signature).svg
}}

'''Barbara McClintock''' (June 16, 1902 – September 2, 1992) was an American scientist and [[cytogenetics|cytogeneticist]] who was awarded the 1983 [[Nobel Prize in Physiology or Medicine]]. McClintock received her PhD in [[botany]] from [[Cornell University]] in 1927. There she started her career as the leader of the development of [[Maize|maize]] cytogenetics, the focus of her research for the rest of her life. From the late 1920s, McClintock studied [[chromosome]]s and how they change during reproduction in maize. She developed the technique for visualizing maize chromosomes and used microscopic analysis to demonstrate many fundamental genetic ideas. One of those ideas was the notion of [[genetic recombination]] by [[chromosomal crossover|crossing-over]] during [[meiosis]]—a mechanism by which chromosomes exchange information. She is often erroneously credited with producing the first [[genetic linkage|genetic map]] for maize, linking regions of the chromosome to physical traits. She demonstrated the role of the [[telomere]] and [[centromere]], regions of the chromosome that are important in the conservation of [[genetic information]]. She was recognized as among the best in the field, awarded prestigious fellowships, and elected a member of the [[National Academy of Sciences]] in 1944.

During the 1940s and 1950s, McClintock discovered [[transposon]]s and used it to demonstrate that [[gene]]s are responsible for turning physical characteristics on and off. She developed theories to explain the suppression and expression of genetic information from one generation of maize plants to the next. Due to skepticism of her research and its implications, she stopped publishing her data in 1953.

Later, she made an extensive study of the cytogenetics and [[ethnobotany]] of maize [[race (biology)|race]]s from South America. McClintock's research became well understood in the 1960s and 1970s, as other scientists confirmed the mechanisms of genetic change and [[regulation of gene expression|protein expression]] that she had demonstrated in her maize research in the 1940s and 1950s. Awards and recognition for her contributions to the field followed, including the Nobel Prize in Physiology or Medicine, awarded to her in 1983 for the discovery of [[Transposable element|genetic transposition]]; as of 2025, she remains the only woman who has received an unshared Nobel Prize in that category.{{sfn|Nobel Prize}}

==Early life==
[[File:McClintock family 1907.jpg|thumb|300px|McClintock children, from left to right: Mignon, Malcolm Rider "Tom", Barbara and Marjorie]]
[[File:McClintock family.jpg|thumb|300px|McClintock family, from left to right: Mignon, Tom, Barbara, Marjorie and Sara (at the piano)]]
Barbara McClintock was born '''Eleanor McClintock''' on June 16, 1902, in [[Hartford, Connecticut|Hartford]], Connecticut,{{sfn|Lamberts|2000}}{{sfn|Comfort|2001|pp=19–22}} the third of four children born to homeopathic physician Thomas Henry McClintock and Sara Handy McClintock.<ref>{{Cite book|title=American women scientists: 23 inspiring biographies, 1900–2000| last=Reynolds | first=Moira Davison |date=1999|publisher=McFarland|isbn=978-0-7864-0649-4|location=Jefferson, N.C.|oclc=40926627}}</ref> Thomas McClintock was the child of British immigrants. Marjorie, the oldest child, was born in October 1898; Mignon, the second daughter, was born in November 1900. The youngest, Malcolm Rider (called Tom), was born 18 months after Barbara. When she was a young girl, her parents determined that [[Eleanor]], a "feminine" and "delicate" name, was not appropriate for her, and chose [[Barbara (given name)|Barbara]] instead.{{sfn|Comfort|2001|pp=19–22}}{{sfn|Keller|1983}}  McClintock was an independent child beginning at a very young age, a trait she later identified as her "capacity to be alone". From the age of three until she began school, McClintock lived with an aunt and uncle in Brooklyn, New York, in order to reduce the financial burden on her parents while her father established his medical practice. She was described as a solitary and independent child. She was close to her father, but had a difficult relationship with her mother, tension that began when she was young.{{sfn|Comfort|2001|pp=19–22}}{{sfn|Keller|1983}}

The McClintock family moved to Brooklyn in 1908 and McClintock completed her secondary education there at [[Erasmus Hall High School]];{{sfn|Keller|1983}}{{sfn|Boyer|2001}} she graduated in 1919.{{sfn|Kass|2024|p=11}} She discovered her love of science and reaffirmed her solitary personality during high school.{{sfn|Comfort|2001|pp=19–22}} She wanted to continue her studies at [[Cornell University]]'s College of Agriculture. Her mother resisted sending McClintock to college for fear that she would be unmarriageable, a common attitude at the time.{{sfn|Keller|1983}} McClintock was almost prevented from starting college, but her father allowed her to just before registration began, and she matriculated at Cornell in 1919.{{sfn|Comfort|2001|pp=23–27}}{{sfn|Fedoroff|1995|p=215}}

==Education and research at Cornell==
{{Listen|type=speech|pos=right|filename=Barbara McClintock (As Told By Jo Handelsman).ogg|title=Barbara McClintock (As Told By Jo Handelsman)|description= }}
McClintock began her studies at Cornell's [[Cornell University College of Agriculture and Life Sciences|College of Agriculture]] in 1919. There, she participated in [[student government]] and was invited to join a [[sorority]], but later broke her sorority pledge.{{sfn|Kass|2024|p=18}} Instead, McClintock took up music, specifically [[jazz]]. She studied [[botany]], receiving a [[Bachelor of Science|BSc]] in 1923.{{sfn|Comfort|2001|pp=23–27}} Her interest in genetics began when she took her first course in that field in 1921. The course was based on a similar one offered at Harvard University, and was taught by [[C. B. Hutchison]], a plant breeder and geneticist.{{sfn|Kass|Provine|1997|p=123}}{{sfn|Kass|2000|p=64}}{{sfn|Fedoroff|1995|p=216}} Hutchison was impressed by McClintock's interest, and telephoned to invite her to participate in the graduate genetics course at Cornell in 1922. McClintock pointed to Hutchison's invitation as a catalyst for her interest in genetics: "Obviously, this telephone call cast the die for my future. I remained with genetics thereafter."{{sfn|McClintock|1983}} Although it has been reported that women could not major in genetics at Cornell, and therefore her MS and PhD—earned in 1925 and 1927, respectively—were officially awarded in botany, recent research has revealed that women were permitted to earn graduate degrees in Cornell's Plant Breeding Department during the time that McClintock was a student at Cornell.{{sfn|Kass|2003|pp=1251–1260}}{{sfn|Kass|2024|pp=14–27}}

During her graduate studies and postgraduate appointment as a botany instructor, McClintock was instrumental in assembling a group that studied the new field of [[cytogenetics]] in maize. This group brought together plant breeders and cytologists, and included [[Marcus Rhoades]], future [[Nobel Prize|Nobel]] laureate [[George Beadle]], and [[Harriet Creighton]].{{sfn|Kass|2005a|pp=118–125}}{{sfn|Kass|2007}}{{sfn|The Barbara McClintock Papers – Cornell}} [[Rollins A. Emerson]], head of the Plant Breeding Department, supported these efforts, although he was not a cytologist himself.{{sfn|Kass|Bonneuil|2004|pp=91–118}}{{sfn|Kass|Bonneuil|Coe|2005}}{{sfn|Kass|2024|pp=46–66}}

She also worked as a research assistant for [[Lowell Fitz Randolph]] and then for [[Lester W. Sharp]], both Cornell botanists.<ref>{{cite web|first=Federica Turriziani |last=Colonna |title=Barbara McClintock (1902–1992) |url=http://embryo.asu.edu/pages/barbara-mcclintock-1902-1992 |website=Embryo.asu.edu |access-date=November 27, 2014}}</ref>{{sfn|Kass|2024|pp=30–42}}

McClintock's cytogenetic research focused on developing ways to visualize and characterize maize chromosomes. This particular part of her work influenced a generation of students, as it was included in most textbooks. She also developed a technique using [[carmine]] staining to visualize maize chromosomes, and showed for the first time the morphology of the 10 maize chromosomes. This discovery was made because she observed cells from the [[microspore]] as opposed to the [[root cap|root tip]].{{sfn|Kass|Bonneuil|2004|pp=91–118}}{{sfn|Fedoroff|1995|p=217}} {{sfn|Kass|2024|pp=46–66}} By studying the morphology of the chromosomes, McClintock was able to [[genetic linkage|link]] specific chromosome groups of traits that were inherited together.{{sfn|Fedoroff|1995|p=212}} Marcus Rhoades noted that McClintock's 1929 ''[[Genetics (journal)|Genetics]]'' paper on the characterization of [[polyploidy|triploid]] maize chromosomes triggered scientific interest in maize cytogenetics, and attributed to her 10 of the 17 significant advances in the field that were made by Cornell scientists between 1929 and 1935.{{sfn|Rhoades}}

In 1930, McClintock was the first person to describe the cross-shaped interaction of homologous chromosomes during [[meiosis]]. The following year, McClintock and Creighton proved the link between [[chromosomal crossover]] during meiosis and the recombination of genetic traits.{{sfn|Fedoroff|1995|p=212}}{{sfn|Coe|Kass|2005|pp=6641–6656}}{{sfn|Kass|2024|pp=75–79}} They observed how the recombination of chromosomes seen under a microscope correlated with new traits.{{sfn|The Barbara McClintock Papers – Cornell}}{{sfn|Creighton|McClintock|1931|pp=492–497}} Until this point, it had only been hypothesized that [[genetic recombination]] could occur during meiosis, although it had not been shown genetically.{{sfn|The Barbara McClintock Papers – Cornell}} It is often reported that McClintock published the first genetic map for maize in 1931, showing the order of three genes on maize chromosome 9;{{sfn|McClintock|1931|pp=485–491}} however, it was her genetics professor C. B. Hutchison who has published the first genetic linkage maps for Chromosome 9 in 1921 and 1922.{{sfn|Kass|2024|p=50}} Her chromosome map showed that the arrangement of genes was consistent with the linkage map published by Hutchison in 1921.{{sfn|Kass|2024|p=51}} This information provided necessary data for the crossing-over study she published with Creighton;{{sfn|Coe|Kass|2005|pp=6641–6656}} they also showed that crossing-over occurs in [[sister chromatids]] as well as [[homologous chromosomes]].{{sfn|Fedoroff|1995|p=218}} In 1938, she produced a cytogenetic analysis of the [[centromere]], describing the organization and function of the centromere, as well as the fact that it can divide.{{sfn|Fedoroff|1995|p=212}}

McClintock's breakthrough publications, and support from her colleagues, led to her being awarded several postdoctoral fellowships from the [[United States National Research Council|National Research Council]]. This funding allowed her to continue to study genetics at Cornell, the [[University of Missouri]], and the [[California Institute of Technology]], where she worked with E. G. Anderson.{{sfn|Kass|2003|pp=1251–1260}}{{sfn|Fedoroff|1995|p=218}} During the summers of 1931 and 1932, she worked at the University of Missouri with geneticist [[Lewis Stadler]], who introduced her to the use of [[X-ray]]s as a [[mutagen]]. Exposure to X-rays can increase the rate of mutation above the natural background level, making it a powerful research tool for genetics. Through her work with X-ray-mutagenized maize, she identified [[ring chromosome]]s, which form when the ends of a single chromosome fuse together after radiation damage.{{sfn|Fedoroff|1995|p=219}}  McClintock found the first ring chromosome in maize in 1931, but the first ring chromosome was first reported by Michael Navashin, who she cited in her first study with Stadler.{{sfn|Kass|2024|p=79}} From this evidence, McClintock hypothesized that there must be a structure on the chromosome tip that would normally ensure stability. She showed that the loss of ring-chromosomes at meiosis caused [[variegation]] in maize foliage in generations subsequent to irradiation resulting from chromosomal deletion.{{sfn|Fedoroff|1995|p=212}} During this period, she demonstrated the presence of the [[nucleolus organizer region]] on a region on maize chromosome 6, which is required for the assembly of the [[nucleolus]].{{sfn|Fedoroff|1995|p=212}}{{sfn|Fedoroff|1995|p=218}}{{sfn|McClintock|1934|p=294–328}} In 1933, she established that cells can be damaged when [[Non-homologous end joining|nonhomologous recombination]] occurs.{{sfn|Fedoroff|1995|p=212}}{{sfn|Green|1959|p=1243}} During this same period, McClintock hypothesized that the tips of chromosomes are protected by [[telomere]]s.{{sfn|CSHL Biography}}

McClintock received a fellowship from the [[John Simon Guggenheim Memorial Foundation|Guggenheim Foundation]] that made possible six months of training in Germany during 1933 and 1934.{{sfn|Fedoroff|1995|p=219}} She had planned to work with [[Curt Stern (geneticist)|Curt Stern]], who had demonstrated crossing-over in ''[[Drosophila]]'' just weeks after McClintock and Creighton had done so; however, Stern emigrated to the United States. Instead,  she worked with geneticist [[Richard B. Goldschmidt]], who was a director of the Kaiser Wilhelm Institute for Biology in Berlin.{{sfn|Keller|1983}}<ref>{{cite journal |last1=Dietrich |first1=M.R. |s2cid=13478874 |title=Richard Goldschmidt: hopeful monsters and other 'heresies' |journal=Nature Reviews Genetics |date=2003 |volume=4 |issue=1 |pages=68–74 |doi=10.1038/nrg979|pmid=12509755 }}</ref> She left Germany early amidst mounting political tension in Europe, returned to Cornell, but found that the university would not hire a woman professor.<ref>{{Cite magazine |last1=Magazine |first1=Smithsonian |last2=Doudna |first2=Jennifer |title=By Studying Corn, Barbara McClintock Unlocked the Secrets of Life |url=https://www.smithsonianmag.com/smithsonian-institution/by-studying-corn-barbara-mcclintock-unlocked-secrets-life-180981555/ |access-date=February 14, 2023 |magazine=Smithsonian Magazine |language=en}}</ref> Legends such as the one cited here are based on undocumented evidence. Kass 2024 has evidence that McClintock worked at Cornell on her return to Cornell after spending only 5 months in Germany.  Emerson hired her as his assistant in the Department of Plant Breeding where McClintock conducted independent research which led to a job offer as an assistant professor at the University of Missouri.{{sfn|Kass|2024|pp=92–95}} In 1936, she accepted an Assistant Professorship offered to her by Lewis Stadler in the Department of Botany at the University of Missouri in [[Columbia, Missouri|Columbia]].{{sfn|Kass|2005b|p=52–71}}{{sfn|Fedoroff|1995|p=220}} While still at Cornell, she was supported by a two-year [[Rockefeller Foundation]] grant obtained for her through Emerson's efforts.{{sfn|Fedoroff|1995|p=219}}

==University of Missouri==
During her time at Missouri, McClintock expanded her research on the effect of X-rays on maize cytogenetics. McClintock observed the breakage and fusion of chromosomes in irradiated maize cells. She was also able to show that, in some plants, spontaneous chromosome breakage occurred in the cells of the endosperm. Over the course of [[mitosis]], she observed that the ends of broken chromatids were rejoined after the chromosome [[DNA replication|replication]].{{sfn|The Barbara McClintock Papers – Missouri}} In the [[anaphase]] of mitosis, the broken chromosomes formed a chromatid bridge, which was broken when the chromatids moved towards the cell poles. The broken ends were rejoined in the [[interphase]] of the next mitosis, and the cycle was repeated, causing massive mutation, which she could detect as variegation in the endosperm.{{sfn|McClintock|1941|pp=234–282}} This [[Breakage-fusion-bridge cycle|breakage–rejoining–bridge cycle]] was a key cytogenetic discovery for several reasons.{{sfn|The Barbara McClintock Papers – Missouri}} First, it showed that the rejoining of chromosomes was not a random event, and second, it demonstrated a source of large-scale mutation. For this reason, it remains an area of interest in cancer research today.{{sfn|Selvarajah et al. 2006}}

Although her research was progressing at Missouri, McClintock was not satisfied with her position at the university. She recalled being excluded from faculty meetings, and was not made aware of positions available at other institutions.{{sfn|Keller|1983}} In 1940, she wrote to Charles Burnham, "I have decided that I must look for another job. As far as I can make out, there is nothing more for me here. I am an assistant professor at $3,000 and I feel sure that that is the limit for me."{{sfn|McClintock|1940}}{{sfn|The Barbara McClintock Papers – Missouri}} Initially, McClintock's position was created especially for her by Stadler, and might have depended on his presence at the university.{{sfn|Kass|2003|pp=1251–1260}}{{sfn|Kass|2005b|p=52–71}} McClintock believed she would not gain [[tenure]] at Missouri, even though according to some accounts, she knew she would be offered a promotion from Missouri in the spring of 1942.{{sfn|Comfort|2002|p=440}} Recent evidence reveals that McClintock more likely decided to leave Missouri because she had lost trust in her employer and in the university administration, after discovering that her job would be in jeopardy if Stadler were to leave for [[Caltech]], as he had considered doing. The university's retaliation against Stadler amplified her sentiments.{{sfn|Kass|2003|pp=1251–1260}}{{sfn|Kass|2024|pp=100–131}}

In early 1941, she took a leave of absence from Missouri in hopes of finding a position elsewhere. She accepted a visiting professorship at [[Columbia University]], where her former Cornell colleague Marcus Rhoades was a professor. Rhoades also offered to share his research field at Cold Spring Harbor on Long Island. In December 1941, she was offered a temporary research position by [[Milislav Demerec]], the newly appointed acting director of the [[Carnegie Institution of Washington]]'s Department of Genetics at [[Cold Spring Harbor Laboratory]]. Although hesitant to make a long-term commitment, McClintock accepted the invitation and became a permanent member of the staff in 1943.{{sfn|Fedoroff|1995|p=221}}{{sfn|Kass|2024|p=128}}

==Cold Spring Harbor==
After her year-long temporary appointment, McClintock accepted a full-time research position at [[Cold Spring Harbor Laboratory]]. There, she was highly productive and continued her work with the breakage-fusion-bridge cycle, using it to substitute for X-rays as a tool for mapping new genes. In 1944, in recognition of her prominence in the field of genetics during this period, McClintock was elected to the [[United States National Academy of Sciences|National Academy of Sciences]]—only the third woman to be elected. The following year she became the first female president of the [[Genetics Society of America]];{{sfn|Lamberts|2000}} she had been elected its vice-president in 1939.{{sfn|Fedoroff|1995|p=220}} In 1944 she undertook a cytogenetic analysis of ''[[Neurospora crassa]]'' at the suggestion of George Beadle, who used the fungus to demonstrate the one gene–one enzyme relationship. He invited her to [[Stanford University|Stanford]] to undertake the study. She successfully described the number of chromosomes, or [[karyotype]], of ''N. crassa'' and described the entire life cycle of the species. Beadle said, "Barbara, in two months at Stanford, did more to clean up the cytology of ''Neurospora'' than all other cytological geneticists had done in all previous time on all forms of mold."{{sfn|Keller|1983|p=114}} ''N. crassa'' has since become a [[model organism|model species]] for classical genetic analysis.{{sfn|McClintock|1945|pp=671–678}}{{sfn|Klug|Cummings|Spencer|Palladino|2012|pp=128–130}}

===Discovery of controlling elements===
[[File:Corn mosaic.jpg|thumb|The relationship of ''Ac/Ds'' in the control of the elements and mosaic color of maize. The seed in 10 is colorless; there is no ''Ac'' element present, and ''Ds'' inhibits the synthesis of colored pigments called [[anthocyanin]]s. In 11 to 13, one copy of ''Ac'' is present; ''Ds'' can move, and some anthocyanin is produced, creating a mosaic pattern. In the kernel in panel 14 there are two ''Ac'' elements, and in panel 15 there are three.<!-- FAIR USE of File:Corn mosaic.jpg: see image description page at http://en.wikipedia.org/wiki/File:Corn mosaic.jpg for rationale -->]]

In the summer of 1944 at Cold Spring Harbor Laboratory, McClintock began systematic studies on the mechanisms of the [[mosaic (genetics)|mosaic]] color patterns of maize seed and the unstable [[Biological inheritance|inheritance]] of this mosaicism.{{sfn|Comfort|2001|pp=84–94}} She identified two new [[dominance relationship|dominant]] and interacting genetic loci that she named ''Dissociation'' (''Ds'') and ''Activator'' (''Ac''). She found that the ''Dissociation'' did not just dissociate or cause the chromosome to break, it also had a variety of effects on neighboring genes when the ''Activator'' was also present, which included making certain stable mutations unstable. In early 1948, she made the surprising discovery that both ''Dissociation'' and ''Activator'' could transpose, or change position, on the chromosome.{{sfn|The Barbara McClintock Papers – Cold Spring Harbor}}{{sfn|Comfort|1999|pp=133–162}}{{sfn|Goodier|Kazazian|2008|pp=23–25}}{{sfn|Fedoroff|1995|p=223}}

She observed the effects of the transposition of ''Ac'' and ''Ds'' by the changing patterns of coloration in maize kernels over generations of controlled crosses, and described the relationship between the two [[locus (genetics)|loci]] through intricate microscopic analysis.{{sfn|National Academy of Sciences 2005}}{{sfn|Comfort|2001|pp=102–115}} She concluded that ''Ac'' controls the transposition of the ''Ds'' from chromosome&nbsp;9, and that the movement of ''Ds'' is accompanied by the breakage of the chromosome.{{sfn|Fedoroff|1995|p=223}} When ''Ds'' moves, the [[aleurone]]-color gene is released from the suppressing effect of the ''Ds'' and transformed into the active form, which initiates the pigment synthesis in cells.{{sfn|Klug|Cummings|Spencer|Palladino|2012|p=395}} The transposition of ''Ds'' in different cells is random, it may move in some but not others, which causes color mosaicism. The size of the colored spot on the seed is determined by stage of the seed development during dissociation. McClintock also found that the transposition of ''Ds'' is determined by the number of ''Ac'' copies in the cell.{{sfn|Pray|Zhaurova|2008}}

Between 1948 and 1950, she developed a theory by which these mobile elements regulated the genes by inhibiting or modulating their action. She referred to ''Dissociation'' and ''Activator'' as "controlling units"—later, as "controlling elements"—to distinguish them from genes. She hypothesized that [[gene regulation]] could explain how complex multicellular organisms made of cells with identical [[genome]]s have cells of different function.{{sfn|Pray|Zhaurova|2008}} McClintock's discovery challenged the concept of the genome as a static set of instructions passed between generations.{{sfn|Lamberts|2000}} In 1950, she reported her work on ''Ac/Ds'' and her ideas about gene regulation in a paper entitled "The origin and behavior of mutable loci in maize" published in the journal ''[[Proceedings of the National Academy of Sciences]]''. In summer 1951, she reported her work on the origin and behavior of mutable loci in maize at the annual symposium at Cold Spring Harbor Laboratory, presenting a paper of the same name. The paper delved into the instability caused by ''Ds'' and ''Ac'' or just ''Ac'' in four genes, along with the tendency of those genes to unpredictably revert to the wild phenotype. She also identified "families" of transposons, which did not interact with one another.{{sfn|Lamberts|2000}}{{sfn|McClintock|1950|pp=344–355}}{{sfn|Fedoroff|1995|p=223}}

[[File:McClintock.jpg|thumb|McClintock with one of her results]]
Her work on controlling elements and gene regulation was conceptually difficult and was not immediately understood or accepted by her contemporaries; she described the reception of her research as "puzzlement, even hostility".{{sfn|McClintock|1987}}{{sfn|Fedoroff|1995|p=223}} Nevertheless, McClintock continued to develop her ideas on controlling elements. She published a paper in ''Genetics'' in 1953, where she presented all her statistical data, and undertook lecture tours to universities throughout the 1950s to speak about her work.{{sfn|McClintock|1953|pp=579–599}} She continued to investigate the problem and identified a new element that she called ''Suppressor-mutator'' (''Spm''), which, although similar to ''Ac/Ds'', acts in a more complex manner. Like ''Ac/Ds'', some versions could transpose on their own and some could not; unlike ''Ac/Ds'', when present, it fully suppressed the expression of mutant genes when they normally would not be entirely suppressed.{{sfn|Fedoroff|1995|p=224}} Based on the reactions of other scientists to her work, McClintock felt she risked alienating the scientific mainstream, and from 1953 was forced to stop publishing accounts of her research on controlling elements.{{sfn|Lamberts|2000}}{{sfn|Comfort|1999|pp=133–162}}

===The origins of maize===
[[File:Corn and microscope.jpg|thumb|McClintock's microscope and ears of corn on exhibition at the [[National Museum of Natural History]] in Washington, D.C.]]

In 1957, McClintock received funding from the [[National Academy of Sciences]] to start research on indigenous strains of maize in Central America and South America. She was interested in studying the [[evolution]] of maize through chromosomal changes,{{sfn|Fedoroff|1995|p=226}} and being in South America would allow her to work on a larger scale. McClintock explored the chromosomal, morphological, and evolutionary characteristics of various [[race (biology)|races]] of maize.{{sfn|Comfort|2001|pp=209–217}}{{sfn|CSHL Biography}} After extensive work in the 1960s and 1970s, McClintock and her collaborators published the seminal study ''The Chromosomal Constitution of Races of Maize'', leaving their mark on [[paleobotany]], [[ethnobotany]], and [[evolutionary biology]].{{sfn|The Barbara McClintock Papers – Origins of Maize}} See Kass (2024, Chapter 9), for extensive details of McClintock's research and influence on the cytogenetics of the Races of Maize in Mexico and South America.

===Rediscovery===
McClintock officially retired from her position at the Carnegie Institution in 1967,{{sfn|Lamberts|2000}} and was made a Distinguished Service Member of the Carnegie Institution of Washington.{{sfn|Fedoroff|1995|p=221}} This honor allowed her to continue working with graduate students and colleagues in the Cold Spring Harbor Laboratory as ''scientist emerita''; she lived in the town.{{sfn|Kolata|1992}} In reference to her decision 20 years earlier to stop publishing detailed accounts of her work on controlling elements, she wrote in 1973:

{{blockquote|Over the years I have found that it is difficult if not impossible to bring to consciousness of another person the nature of his tacit assumptions when, by some special experiences, I have been made aware of them. This became painfully evident to me in my attempts during the 1950s to convince geneticists that the action of genes had to be and was controlled. It is now equally painful to recognize the fixity of assumptions that many persons hold on the nature of controlling elements in maize and the manners of their operation. One must await the right time for conceptual change.{{sfn|McClintock|1973}}}}

The importance of McClintock's contributions was revealed in the 1960s, when the work of French geneticists [[François Jacob]] and [[Jacques Monod]] described the genetic regulation of the [[lac operon|''lac'' operon]], a concept she had demonstrated with ''Ac/Ds'' in 1951. Following Jacob and Monod's 1961 ''Journal of Molecular Biology'' paper "Genetic regulatory mechanisms in the synthesis of proteins", McClintock wrote an article for ''[[American Naturalist]]'' comparing the ''lac'' operon and her work on controlling elements in maize.{{sfn|McClintock|1961|pp=265–277}}{{sfn|Fedoroff|1995|p=224}} Even late in the twentieth century, McClintock's contribution to biology was still not widely acknowledged as amounting to the discovery of genetic regulation.{{sfn|Comfort|1999|pp=133–162}} See Kass (2024, pp.&nbsp;189–191) for clarification of this legend.

McClintock was widely credited with discovering transposition after other researchers finally discovered the process in bacteria, yeast, and [[bacteriophage]]s in the late 1960s and early 1970s.{{sfn|Fedoroff|1995|p=213}} During this period, molecular biology had developed significant new technology, and scientists were able to show the molecular basis for transposition.{{sfn|Fedoroff|1995|p=227}} In the 1970s, ''Ac'' and ''Ds'' were [[cloning|cloned]] by other scientists and were shown to be [[transposon|class II transposons]]. ''Ac'' is a complete transposon that can produce a functional [[transposase]], which is required for the element to move within the genome. ''Ds'' has a mutation in its transposase gene, which means that it cannot move without another source of transposase. Thus, as McClintock observed, ''Ds'' cannot move in the absence of ''Ac''.<ref>{{cite journal |url=https://www.nature.com/scitable/topicpage/barbara-mcclintock-and-the-discovery-of-jumping-34083/|title=Barbara McClintock and the Discovery of Jumping Genes (Transposons)|last1=Pray |first1=Leslie |last2=Zhaurova |first2=Kira |journal=Nature Education |volume=1 |issue=1 |page=169}}</ref> ''Spm'' has also been characterized as a transposon. Subsequent research has shown that transposons typically do not move unless the cell is placed under stress, such as by irradiation or the breakage-fusion-bridge cycle, and thus their activation during stress can serve as a source of genetic variation for evolution.{{sfn|Pray|2008}} McClintock understood the role of transposons in evolution and genome change well before other researchers grasped the concept. Nowadays, ''Ac/Ds'' is used as a tool in plant biology to generate mutant plants used for the characterization of gene function.{{sfn|Jin|Duan|Zhang|Chen|2003}}

==Honors and recognition==
In 1947, McClintock received the Achievement Award from the [[American Association of University Women]]. She was elected a Fellow of the [[American Academy of Arts and Sciences]] in 1959.<ref name=AAAS>{{cite web|title=Book of Members, 1780–2010: Chapter M|url=http://www.amacad.org/publications/BookofMembers/ChapterM.pdf|publisher=American Academy of Arts and Sciences|access-date=July 22, 2014}}</ref> In 1967, McClintock was awarded the Kimber Genetics Award;{{sfn|Fedoroff|1995|p=229}} three years later, she was given the [[National Medal of Science]] by [[Richard Nixon]] in 1970.{{sfn|Fedoroff|1995|p=213}}{{sfn|National Medal of Science}} She was the first woman to be awarded the National Medal of Science.<ref>{{cite web |url=https://www.nsf.gov/od/nms/recip_details.jsp?recip_id=234|title=The President's National Medal of Science: Recipient Detailswork=National Science Foundation|archive-url=https://web.archive.org/web/20150911174454/https://www.nsf.gov/od/nms/recip_details.jsp?recip_id=234|archive-date=September 11, 2015}}</ref>
Cold Spring Harbor named a building in her honor in 1973.{{sfn|CSHL Biography}} She received the Louis and Bert Freedman Foundation Award and the Lewis S. Rosensteil Award in 1978.{{sfn|Fedoroff|1995|p=229}} In 1981, she became the first recipient of the [[MacArthur Foundation]] Grant, and was awarded the [[Albert Lasker Award for Basic Medical Research]],{{sfn|Washington Post}} the [[Wolf Prize in Medicine]] and the [[Thomas Hunt Morgan Medal]] by the Genetics Society of America. In 1982, she was awarded the [[Louisa Gross Horwitz Prize]] from [[Columbia University]] for her research in the "evolution of genetic information and the control of its expression."{{sfn|Louisa Gross Horwitz Prize}}{{sfn|CSHL Biography}}

Most notably, she received the [[Nobel Prize for Physiology or Medicine]] in 1983, the first woman to win that prize unshared,{{sfn|Kolata|1992}} and the first American woman to win any unshared Nobel Prize in the sciences.<ref>{{cite web|url=http://www.nasonline.org/member-directory/deceased-members/52803.html |title=Barbara McClintock |website=Nasonline.org |date=March 30, 2018 |access-date=August 19, 2018}}</ref> It was given to her by the [[Nobel Foundation]] for discovering "[[transposon|mobile genetic elements]]";{{sfn|Nobel Prize 1983}} this was more than 30 years after she initially described the phenomenon of controlling elements. She was compared to [[Gregor Mendel]] in terms of her scientific career by the [[Swedish Academy of Sciences]] when she was awarded the Prize.{{sfn|Keirns 1999}} Nobel Prize awards and their limitations, and the rationale for delayed awards are detailed in Kass (2024, pp.&nbsp;236–247).

[[File:McClintock Nobel Lecture.jpg|thumb|McClintock giving her Nobel Lecture]]

She was elected a [[List of fellows of the Royal Society elected in 1989|Foreign Member of the Royal Society (ForMemRS) in 1989]].{{sfn|Fedoroff|1995|p=236}} McClintock received the [[Benjamin Franklin Medal for Distinguished Achievement in the Sciences]] of the [[American Philosophical Society]] in 1993.{{sfn|Benjamin Franklin Medal}} She had been previously elected to the APS in 1946.<ref>{{Cite web|title=APS Member History|url=https://search.amphilsoc.org/memhist/search?creator=Barbara+mcclintock&title=&subject=&subdiv=&mem=&year=&year-max=&dead=&keyword=&smode=advanced|access-date=May 3, 2021|website=search.amphilsoc.org}}</ref> She was awarded 14 Honorary Doctor of Science degrees and an Honorary Doctor of Humane Letters.{{sfn|CSHL Biography}} In 1986 she was inducted into the [[National Women's Hall of Fame]]. During her final years, McClintock led a more public life, especially after [[Evelyn Fox Keller]]'s 1983 biography of her, ''A Feeling for the Organism,'' brought McClintock's story to the public. She remained a regular presence in the Cold Spring Harbor community, and gave talks on mobile genetic elements and the history of genetics research for the benefit of junior scientists. An anthology of her 43 publications ''The Discovery and Characterization of Transposable Elements: The Collected Papers of Barbara McClintock'' was published in 1987.{{sfn|Washington Post}}

The [[McClintock Prize]] is named in her honor.<ref name=mcprize>{{cite web|url=https://www.maizegdb.org/community/awards/mcclintock|title=The McClintock Prize for Plant Genetics and Genome Studies|website=Maizegdb.org}}</ref> Laureates of the award include [[David Baulcombe]], [[Detlef Weigel]], [[Robert A. Martienssen]], [[Jeffrey D. Palmer]]  and [[Susan R. Wessler]].<ref name=mcprize/>

In May 2005 the U.S. Postal Service issued a panel of first-class stamps honoring Barbara McClintock, along with [[Richard Feynman]], [[Josiah Willard Gibbs]], and [[John von Neumann]].<ref>{{cite journal|doi=10.1063/1.1995738 |title=US Postage Stamps Feature Scientists |year=2005 |last1=Feder |first1=Toni |journal=Physics Today |volume=58 |issue=5 |page=30 |bibcode=2005PhT....58e..30F |url=https://pubs.aip.org/physicstoday/article-abstract/58/5/30/1016907/US-Postage-Stamps-Feature-Scientists?redirectedFrom=fulltext|url-access=subscription }}</ref>

In May 2024, a plant species, ''Stellaria mcclintockiae,'' was named after her.<ref>{{Cite news |date=2024-05-26 |title=New plant species from Kerala's Nelliyampathy named after Nobel laureate Barbara McClintock |url=https://www.thehindu.com/news/national/kerala/new-plant-species-from-keralas-nelliyampathy-named-after-nobel-laureate-barbara-mcclintock/article68218141.ece |access-date=2025-03-07 |work=The Hindu |language=en-IN |issn=0971-751X}}</ref>

==Later years==
McClintock spent her later years, post Nobel Prize, as a key leader and researcher in the field at Cold Spring Harbor Laboratory on Long Island, New York.  McClintock died of natural causes in [[Huntington, New York|Huntington]], New York, on September 2, 1992, at the age of 90; she never married or had children.{{sfn|Kolata|1992}}{{sfn|Washington Post}}<ref name="Campbell">{{cite journal |last1=Campbell |first1=Allan |title=Barbara McClintock |journal=Annual Review of Genetics |date=December 1993 |volume=27 |issue=1 |pages=1–32 |doi=10.1146/annurev.ge.27.120193.000245 |pmid=8122898 |language=en |issn=0066-4197|doi-access=free }}</ref>

==Legacy==
McClintock was the subject of a 1983 biography by physicist [[Evelyn Fox Keller]], titled ''A Feeling for the Organism''. Keller argued that because McClintock felt like an outsider within her field, (in part, because of her sex) she was able to look at her scientific subjects from a perspective different from the dominant one, leading to several important insights.<ref>{{Cite book|title=A feeling for the organism: the life and work of Barbara McClintock|last1=Keller|first1=Evelyn Fox|year=1993|orig-date=1983|publisher=W.H. Freeman|isbn=0-8050-7458-9|edition=10th anniversary|location=New York|oclc=29715247}}</ref> Keller shows how this led many of her colleagues to reject her ideas and undermine her abilities for many years. For example, when McClintock presented her findings that the genetics of maize did not conform to Mendelian distributions, geneticist [[Sewall Wright]] expressed the belief that she did not understand the underlying mathematics of her work, a belief he had also expressed towards other women at the time.{{sfn|Esther Lederberg}} In addition, geneticist [[Charlotte Auerbach|Lotte Auerbach]] recounted that [[Joshua Lederberg]] returned from a visit to McClintock's lab with the remark: 'By God, that woman is either crazy or a genius.' " As Auerbach recounts, McClintock had thrown Lederberg and his colleagues out after half an hour 'because of their arrogance. She was intolerant of arrogance&nbsp;... She felt she had crossed a desert alone and no one had followed her.'"{{sfn|Keller|1983|p=142}}{{sfn|Esther Lederberg Colleagues}}

In 2001, a second biography, science historian [[Nathaniel C. Comfort]]'s ''The Tangled Field: Barbara McClintock's Search for the Patterns of Genetic Control'' challenged this narrative. Comfort's biography contests the claim that McClintock was marginalized by other scientists, which he calls the "McClintock Myth" and argues was perpetuated both by McClintock herself as well as in the earlier biography by Keller. Comfort, however, asserts that McClintock was not discriminated against because of her gender, citing that she was well regarded by her professional peers, even in the early years of her career.{{sfn|Comfort|1999}}

In 2024, ''From Chromosomes to Mobile Genetic Elements: The Life and Work of Nobel Laureate Barbara McClintock'', a biography by Lee B. Kass, was released.{{sfn|Kass|2024}}

Many recent biographical works on women in science feature accounts of McClintock's work and experience. She is held up as a role model for girls in such works of children's literature as Edith Hope Fine's ''Barbara McClintock, Nobel Prize Geneticist'', Deborah Heiligman's ''Barbara McClintock: Alone in Her Field'' and Mary Kittredge's ''Barbara McClintock''. A recent biography for young adults by Naomi Pasachoff, ''Barbara McClintock, Genius of Genetics'', provides a new perspective, based on the current literature.{{sfn|Pasachoff|2006}}

On May 4, 2005, the [[United States Postal Service]] issued the "American Scientists" commemorative postage stamp series, a set of four 37-cent self-adhesive stamps in several configurations.  The scientists depicted were Barbara McClintock, [[John von Neumann]], [[Josiah Willard Gibbs]], and [[Richard Feynman]]. McClintock was also featured in a 1989 four-stamp issue from Sweden which illustrated the work of eight Nobel Prize-winning geneticists. A laboratory building at Cold Spring Harbor Laboratory was named for her. A street has been named after her in the new "[[Adlershof#Development|Adlershof Development Society]]" science park in [[Berlin]].{{sfn|Berlin}} 
[[File:Barbara McClintock Hall, Cornell University.jpg|thumb|right|upright=.8|Barbara McClintock Hall at [[Cornell University]]]]
A 103,835 square-foot residence hall at [[Cornell University]] was named for McClintock in 2022.<ref>{{Cite web |title=Barbara McClintock Hall |url=https://scl.cornell.edu/residential-life/housing/campus-housing/first-year-undergraduates/residence-halls/barbara-mcclintock-hall |access-date=October 24, 2022 |website=Student & Campus Life {{!}} Cornell University |language=en}}</ref><ref>{{cite web |title=3225-Barbara McClintock Hall Facility Information |url=https://www.fs.cornell.edu/fs/facinfo/fs_facilinfo.cfm?facil_cd=3225 |website=Facilities and Campus Services |access-date=June 27, 2023}}</ref>

Some of McClintock's personality and scientific achievements were referred to in [[Jeffrey Eugenides]]'s 2011 novel ''[[The Marriage Plot]],'' which tells the story of a [[yeast]] geneticist named Leonard who has [[bipolar disorder]]. He works at a laboratory loosely based on Cold Spring Harbor. The character reminiscent of McClintock is a reclusive geneticist at the fictional laboratory, who makes the same discoveries as her factual counterpart.{{sfn|Kolata|2012}}

Judith Pratt wrote a play about McClintock, called MAIZE, which was read at Artemesia Theatre in Chicago in 2015, and was produced in Ithaca NY, the home of Cornell University, in February–March 2018.<ref>{{cite news |title=MAIZE, a play about Barbara McClintock |url=https://events.cornell.edu/event/maize_a_play_about_barbara_mcclintock |access-date=November 21, 2019 |work=Cornell University |date=March 24, 2018 |language=en}}</ref>
{{clear}}

McClintock’s life continue to inspire novels and essays that reinterpret her life through imaginative storytelling, reflecting her enduring influence on both science and culture.,<ref>{{cite journal |last=Vrettos |first=Athena |title=Genetic Legacies: Fictionalizing Barbara McClintock |journal=Configurations |volume=29 |issue=2 |pages=245–263 |year=2021 |doi=10.1353/con.2021.0012}}</ref> including [[Rachel Pastan]]’s novel ''In the Field'' (2021), which the author describes as an imaginative re-creation of the scientist’s solitude, dedication, and unconventional career.<ref>{{cite web |last=Pastan |first=Rachel |title=So You Want to Write a Novel About a Legendary Corn Geneticist |url=https://lithub.com/so-you-want-to-write-a-novel-about-a-legendary-corn-geneticist/ |website=Literary Hub |publisher=Grove Atlantic and Electric Literature |date=August 12, 2021 |access-date=October 23, 2025}}</ref>

==Key publications==
* {{Cite journal 
| last1 = McClintock | first1 = B. 
| title = A Cytological and Genetical Study of Triploid Maize 
| journal = Genetics 
| volume = 14 
| issue = 2 
| pages = 180–222 
| year = 1929 
| doi = 10.1093/genetics/14.2.180 
| pmid = 17246573 
| pmc = 1201029
 |ref=none}}
* {{Cite journal 
| last1 = Creighton | first1 = H. B. 
| last2 = McClintock | first2 = B. 
| title = A Correlation of Cytological and Genetical Crossing-Over in Zea Mays 
| journal = Proceedings of the National Academy of Sciences of the United States of America 
| volume = 17 
| issue = 8 
| pages = 492–497 
| year = 1931 
| pmid = 16587654 
| pmc = 1076098
 | doi=10.1073/pnas.17.8.492
| bibcode = 1931PNAS...17..492C 
 |ref=none| doi-access = free 
}}
* {{Cite journal 
| last1 = McClintock | first1 = B. 
| title = The Order of the Genes C, Sh and Wx in Zea Mays with Reference to a Cytologically Known Point in the Chromosome 
| journal = Proceedings of the National Academy of Sciences of the United States of America 
| volume = 17 
| issue = 8 
| pages = 485–491 
| year = 1931 
| pmid = 16587653 
| pmc = 1076097
 | doi=10.1073/pnas.17.8.485
| bibcode = 1931PNAS...17..485M 
 |ref=none| doi-access = free 
}}
* {{Cite journal 
| last1 = McClintock | first1 = B. 
| title = The Stability of Broken Ends of Chromosomes in Zea Mays 
| journal = Genetics 
| volume = 26 
| issue = 2 
| pages = 234–282 
| year = 1941 
| doi = 10.1093/genetics/26.2.234 
| pmid = 17247004 
| pmc = 1209127
 |ref=none}}
* {{Cite journal | last1 = McClintock | first1 = B. | title = Neurospora. I. Preliminary Observations of the Chromosomes of Neurospora crassa | journal = American Journal of Botany | volume = 32 | issue = 10 | pages = 671–678 | doi = 10.2307/2437624 | year = 1945 | jstor = 2437624  |ref=none}}
* {{Cite journal 
| doi = 10.1073/pnas.36.6.344 
| last1 = McClintock | first1 = B. 
| title = The origin and behavior of mutable loci in maize 
| journal = Proceedings of the National Academy of Sciences of the United States of America 
| volume = 36 
| issue = 6 
| pages = 344–355 
| year = 1950 
| pmid = 15430309 
| pmc = 1063197
| bibcode = 1950PNAS...36..344M  |ref=none| doi-access = free }}
* {{Cite journal 
| last1 = McClintock | first1 = B. 
| title = Induction of Instability at Selected Loci in Maize 
| journal = Genetics 
| volume = 38 
| issue = 6 
| pages = 579–599 
| year = 1953 
| doi = 10.1093/genetics/38.6.579 
| pmid = 17247459 
| pmc = 1209627
 |ref=none}}
* {{Cite journal | last1 = McClintock | first1 = B. | title = Some Parallels Between Gene Control Systems in Maize and in Bacteria | journal = The American Naturalist | volume = 95 | issue = 884 | pages = 265–277 | doi = 10.1086/282188 | year = 1961 | bibcode = 1961ANat...95..265M | s2cid = 56345866 |ref=none}}
* {{cite book |last1 = McClintock |first1= B. |last2 = Kato Yamakake |first2= T.A. |last3 = Blumenschein |first3= A. |year= 1981 |title= Chromosome constitution of races of maize: Its significance in the interpretation of relationships between races and varieties in the Americas |location=Chapingo, Mexico |publisher=Escuela de Nacional de Agricultura, Colegio de Postgraduados |oclc=9181898 }}
* {{Cite journal | last1 = McClintock | first1 = B. | title = The Significance of Responses of the Genome to Challenge | journal = Science | volume = 226 | issue = 4676 | pages = 792–801 | doi = 10.1126/science.15739260 | year = 1984 | bibcode = 1984Sci...226..792M | pmid = 15739260 |ref=none}}

==See also==
* [[Timeline of women in science]]

==Citations==
{{Reflist|30em}}

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  | year = 2000
  | title = Barbara McClintock: Botanist, cytologist, geneticist. Symposium Botany in the Age of Mendel, Abstract #193
  | journal = American Journal of Botany
  | volume = 87
  | issue = 6
  | page = 64
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  | first = Lee B.
  | year = 2003
  | title = Records and recollections: A new look at Barbara McClintock, Nobel Prize-winning geneticist
  | journal = Genetics
  | issue = 4
  | pages = 1251–1260
  |pmid = 12930736
  | volume = 164
  | doi = 10.1093/genetics/164.4.1251
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* {{citation
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 |title       = Harriet Creighton: Proud botanist
 |journal     = Plant Science Bulletin
 |volume      = 51
 |issue       = 4
 |pages       = 118–125
 |date        = December 2005a
 |url         = http://www.botany.org/PlantScienceBulletin/PSB-2005-51-4.php#HARRIET
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* {{citation
  | last = Kass
  | first = Lee B.
  | date = February 2007
  | title = Harriet B. Creighton (1909–2004), on Women Pioneers in Plant Biology
  | publisher = American Society of Plant Biologists
  | editor = Ann Hirsch
  | url = http://aspb.org/wipb-pioneer-biographies/#tab-id-2
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* {{citation
  | last = Kass
  | first = Lee B.
  | year = 2005b
  | title = Missouri compromise: tenure or freedom. New evidence clarifies why Barbara McClintock left Academe
  | journal = Maize Genetics Cooperation Newsletter
  | issue = 79
  | pages = 52–71
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* {{citation
  | title = Seeing Patterns: Models, Visual Evidence and Pictorial Communication in the Work of Barbara McClintock
  | last = Keirns
  | first = Carla
  | s2cid = 83266829
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  | issue = 1
  | date =Spring 1999
  | pages = 163–196
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* {{citation
  | last = Keller
  | first = Evelyn Fox
  | year = 1983
  | title = A Feeling for the Organism
  | publisher = W. H. Freeman and Company
  | location = New York
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* {{citation
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  | last2 = Cummings
  | first2 = Michael R.
  | last3 = Spencer
  | first3 = Charlotte A.
  | last4 = Palladino
  | first4 = Michael A.
  | title = Concepts of Genetics
  | edition = 10th
  | publisher = Pearson
  | location = Boston
  | isbn = 978-0-321-72412-0
  | year = 2012
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* {{citation
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  | title = Dr. Barbara McClintock, 90, Gene Research Pioneer, Dies
  | work = The New York Times
  | date = September 4, 1992
| last = Kolata
  | first = Gina
  | access-date = December 28, 2012
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* {{citation
  | url = https://www.nytimes.com/2012/02/14/science/yeast-genetics-in-the-marriage-plot-scientist-was-fiction-but-not-his-work.html
  | title = The Scientist Was a Figment, but His Work Was Real
  | work = The New York Times
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  | first = Gina
  | date = February 13, 2012
| access-date = April 5, 2013
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* {{citation
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  | title = Colleagues: Barbara McClintock
  | work = Esther M. Zimmer Lederberg Memorial Website
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| ref ={{sfnRef|Esther Lederberg Colleagues}}
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* {{citation
  | title = The Louisa Gross Horwitz Prize for Biology or Biochemistry
  | url = http://www.cumc.columbia.edu/research/horwitz-prize/prize-awardees
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  | publisher = Columbia University
  | access-date = November 11, 2012
| ref ={{sfnRef|Louisa Gross Horwitz Prize}}
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* {{citation
  | url = http://www.anb.org/articles/13/13-02550.html
  | title = McClintock, Barbara
  | publisher = Oxford University Press
  | work = American National Biography Online
  | last = Lamberts
  | first = William J.
  | date = February 2000
  | access-date = November 28, 2012
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* {{citation
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  | url-status = dead
  | archive-date = June 11, 2011
  | title = Education and Research at Cornell, 1925–1931
  | work = The Barbara McClintock Papers
  | series = Profiles in Science
  | publisher = National Library of Medicine
  | access-date = December 28, 2012
| ref ={{sfnRef|The Barbara McClintock Papers – Cornell}}
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* {{citation
  | title = Breakage-Fusion-Bridge: The University of Missouri, 1936–1941
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  | access-date = November 12, 2012
| ref ={{sfnRef|The Barbara McClintock Papers – Missouri}}
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* {{citation
  | title = Controlling Elements: Cold Spring Harbor, 1942–1967
  | url = https://profiles.nlm.nih.gov/ps/retrieve/Narrative/LL/p-nid/49
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| ref ={{sfnRef|The Barbara McClintock Papers – Cold Spring Harbor}}
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* {{citation
  | ref ={{sfnRef|The Barbara McClintock Papers – Origins of Maize}}
  | title = Searching for the Origins of Maize in South America, 1957–1981
  | work = The Barbara McClintock Papers
  | series = Profiles in Science
  | url = https://profiles.nlm.nih.gov/ps/retrieve/Narrative/LL/p-nid/51
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  | publisher = National Library of Medicine
  | access-date = February 19, 2013
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* {{citation
  | last = McClintock
  | first = Barbara
  | title = A short biographical note: Barbara McClintock
  | year = 1983
  | url = https://www.nobelprize.org/laureate/428
  | publisher = Nobel Foundation
  | access-date = December 28, 2012
}}
* {{citation
  | last = McClintock
  | first = Barbara
  | year = 1931
  | title = The order of the genes C, Sh, and Wx in Zea Mays with reference to a cytologically known point in the chromosome
  | journal = Proceedings of the National Academy of Sciences
  | issue = 8
  | pages = 485–91
  | doi = 10.1073/pnas.17.8.485
  | volume = 17
  | pmid=16587653
  | pmc=1076097
  | bibcode = 1931PNAS...17..485M
  | doi-access = free
 }}
* {{citation
  | last = McClintock
  | first = Barbara
  | s2cid = 24921807
 | title = The relation of a particular chromosomal element to the development of the nucleoli in Zea mays
  | journal = Zeitschrift für Zellforschung und Mikroskopische Anatomie
  | volume = 21
  | issue = 2
  | pages = 294–328
  | year = 1934
  | doi = 10.1007/BF00374060
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | title = The Stability of Broken Ends of Chromosomes in Zea Mays
  | journal = Genetics
  | volume = 26
  | issue = 2
  | pages = 234–82
  |date=March 1941
  | doi = 10.1093/genetics/26.2.234
 | pmid = 17247004
  | pmc = 1209127
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | title = Letter from Barbara McClintock to Charles R. Burnham
  | date = September 16, 1940
| url = https://profiles.nlm.nih.gov/LL/B/B/N/T/_/llbbnt.pdf
  | archive-url = https://web.archive.org/web/20040319142923/http://profiles.nlm.nih.gov/LL/B/B/N/T/_/llbbnt.pdf
  | url-status = dead
  | archive-date = March 19, 2004
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | year = 1945
  | title = ''Neurospora'': preliminary observations of the chromosomes of ''Neurospora crassa''
  | journal = American Journal of Botany
  | volume = 32
  | pages = 671–78
  | jstor = 2437624
  | doi = 10.2307/2437624
  | issue = 10
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | year = 1950
  | title = The origin and behavior of mutable loci in maize
  | journal = Proceedings of the National Academy of Sciences
  | volume = 36
  | pages = 344–55
  | doi = 10.1073/pnas.36.6.344
  | issue = 6
  | pmid=15430309
  | pmc=1063197
  | bibcode = 1950PNAS...36..344M
  | doi-access = free
 }}
* {{citation
  | last = McClintock
  | first = Barbara
  | title = The discovery and characterization of transposable elements: the collected papers of Barbara McClintock
  | editor-first = John A.
  | editor-last = Moore
  | year = 1987
  | publisher = Garland Pub.
  | location = New York
  | isbn = 978-0-8240-1391-2
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | year = 1953
  | title = Induction of instability at selected loci in maize
  | journal = Genetics
  | volume = 38
  | pages = 579–99
  | pmid = 17247459
  | issue = 6
  | doi = 10.1093/genetics/38.6.579
 | pmc = 1209627
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | title = Letter from Barbara McClintock to Maize geneticist Oliver Nelson
  | year = 1973
  | url = https://profiles.nlm.nih.gov/ps/retrieve/Narrative/LL/p-nid/52
  | archive-url = https://web.archive.org/web/20110611105002/http://profiles.nlm.nih.gov/ps/retrieve/Narrative/LL/p-nid/52
  | url-status = dead
  | archive-date = June 11, 2011
  }}
* {{citation
  | last = McClintock
  | first = Barbara
  | year = 1961
  | title = Some parallels between gene control systems in maize and in bacteria
  | journal = American Naturalist
  | volume = 95
  | issue = 884
  | pages = 265–77
  | doi=10.1086/282188
  | bibcode = 1961ANat...95..265M
 | s2cid = 56345866
 }}
* {{citation
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 |ref={{sfnRef|National Academy of Sciences 2005}} 
 |year=2005 
  
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 |archive-date=November 28, 2012 }}
* {{citation
 |url=http://www.nationalmedals.org/medals/1970.php 
 |title=1970 National Medal of Science 
 |publisher=National Science and Technology Medals Foundation 
 |work=The Laureates 
 |year=2009 
 |access-date=December 28, 2012 |ref={{sfnRef|National Medal of Science}} 
  
 |archive-url=https://web.archive.org/web/20120911074221/https://www.nationalmedals.org/medals/1970.php 
 |archive-date=September 11, 2012 }}
* {{citation
  | url = http://nobelprize.org/nobel_prizes/medicine/shortfacts.html
  | title = Facts on the Nobel Prize in Physiology or Medicine
  | work = Nobelprize.org
  | publisher = Nobel Media AB
  | access-date = July 12, 2010
| archive-url = https://web.archive.org/web/20100725010741/http://nobelprize.org/nobel_prizes/medicine/shortfacts.html
  | archive-date = July 25, 2010
| url-status = live
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* {{citation
  | title = The Nobel Prize in Physiology or Medicine 1983
  | url = http://nobelprize.org/nobel_prizes/medicine/laureates/1983/
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  | publisher = Nobel Media AB
  | access-date = July 8, 2010
| archive-url = https://web.archive.org/web/20100706044749/http://nobelprize.org/nobel_prizes/medicine/laureates/1983/
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* {{citation
  | last = Pasachoff
  | first = Naomi
  | year = 2006
  | title = Barbara McClintock, Genius of Genetics
  | publisher = Enslow Publishers, Inc.
  | isbn = 978-0-7660-2505-9
  | url = https://archive.org/details/barbaramcclintoc00pasa
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* {{citation
  | last1 = Pray
  | first1 = Leslie
  | url = http://www.nature.com/scitable/topicpage/transposons-the-jumping-genes-518
  | title = Transposons: The Jumping Genes
  | publisher = Nature Publishing Group
  | journal = Nature Education
  | year = 2008
  | volume = 1
  | issue = 1
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* {{citation
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  | first1 = Leslie
  | last2 = Zhaurova
  | first2 = Kira
  | year = 2008
  | title = Barbara McClintock and the Discovery of Jumping Genes (Transposons)
  | journal = Nature Education
  | volume = 1
  | issue = 1
  | url = http://www.nature.com/scitable/topicpage/barbara-mcclintock-and-the-discovery-of-jumping-34083
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* {{citation
  | last = Rhoades
  | first = Marcus M.
  | title = The golden age of corn genetics at Cornell as seen through the eyes of M. M. Rhoades
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  | archive-url = https://web.archive.org/web/20041108010734/http://profiles.nlm.nih.gov/LL/B/B/F/Z/_/llbbfz.pdf
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* {{citation
  | vauthors = Selvarajah S, Yoshimoto M, Park PC, Maire G, Paderova J, Bayani J, Lim G, Al-Romaih K, Squire JA, Zielenska M
  | s2cid = 12403557
 | title = The breakage-fusion-bridge (BFB) cycle as a mechanism for generating genetic heterogeneity in osteosarcoma
  | journal = Chromosoma
  | date=December 2006
  | volume = 115
  | series = 6
  | pages = 459–467
  | ref ={{sfnRef|Selvarajah et al. 2006}}
  | doi = 10.1007/s00412-006-0074-4
  | issue = 6
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* {{citation
  | url = http://www.highbeam.com/doc/1P2-1023553.html
  | archive-url = https://web.archive.org/web/20141011021333/http://www.highbeam.com/doc/1P2-1023553.html
  
  | archive-date = October 11, 2014
| title = Geneticist B. McClintock Dies; Nobelist
  | newspaper = [[The Washington Post]]
  | date = September 4, 1992
| access-date = December 28, 2012
| ref ={{sfnRef|Washington Post}}
  |url-access=subscription
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{{Refend}}

===Archives and research collections===
* [https://profiles.nlm.nih.gov/spotlight/ll The Barbara McClintock Papers] – Profiles in Science, National Library of Medicine.
* [http://www.amphilsoc.org/mole/view?docId=ead/Mss.Ms.Coll.79-ead.xml;query=;brand=default Barbara McClintock Papers, 1927–1991] at the [[American Philosophical Society]]

==External links==
{{Commons category|Barbara McClintock}}
* {{Wikiquote-inline}}
* [https://web.archive.org/web/20110927171607/http://library.cshl.edu/archives/archives/bmcbio.htm Cold Spring Harbor Laboratory Archives, Barbara McClintock: A Brief Biographical Sketch]
* [http://www.arabidopsis.org/info/springer.jsp Enhancer and Gene Trap Transposon Mutagenesis in Arabidopsis], comprehensive article on the use of ''Ac/Ds'' and other transposons for plant mutagenesis
* [https://www.newscientist.com/people/barbara-mcclintock/ Barbara McClintock] archive on New Scientist
* {{Nobelprize}}
* [https://www.pnas.org/doi/10.1073/pnas.1219372109 Barbara McClintock] on Pnas.org

{{Wolf Prize in Medicine}}
{{Nobel Prize in Physiology or Medicine Laureates 1976-2000}}
{{1983 Nobel Prize winners}}
{{Winners of the National Medal of Science|biological}}
{{Connecticut Women's Hall of Fame}}
{{McClintock Prize Laureates}}
{{National Women's Hall of Fame}}
{{FRS 1989}}
{{Authority control}}

{{Featured article}}

{{DEFAULTSORT:McClintock, Barbara}}
[[Category:American geneticists]]
[[Category:American evolutionary biologists]]
[[Category:American theoretical biologists]]
[[Category:Women Nobel laureates]]
[[Category:1902 births]]
[[Category:1992 deaths]]
[[Category:American women botanists]]
[[Category:American women evolutionary biologists]]
[[Category:Women food scientists]]
[[Category:American women geneticists]]
[[Category:American women physiologists]]
[[Category:American Nobel laureates]]
[[Category:Nobel laureates in Physiology or Medicine]]
[[Category:Fellows of the American Academy of Arts and Sciences]]
[[Category:Members of the United States National Academy of Sciences]]
[[Category:National Medal of Science laureates]]
[[Category:Recipients of the Albert Lasker Award for Basic Medical Research]]
[[Category:Wolf Prize in Medicine laureates]]
[[Category:MacArthur Fellows]]
[[Category:Foreign members of the Royal Society]]
[[Category:Cornell University College of Agriculture and Life Sciences alumni]]
[[Category:Erasmus Hall High School alumni]]
[[Category:University of Missouri faculty]]
[[Category:American people of British descent]]
[[Category:Scientists from Brooklyn]]
[[Category:Scientists from Columbia, Missouri]]
[[Category:Scientists from Hartford, Connecticut]]
[[Category:20th-century American botanists]]
[[Category:20th-century American women scientists]]
[[Category:Scientists from Missouri]]
[[Category:Scientists from New York (state)]]
[[Category:American lecturers]]
[[Category:Plant geneticists]]
[[Category:Graduate Women in Science members]]
[[Category:Members of the American Philosophical Society]]