Lilian Vaughan Morgan (''née'' Sampson; July 7, 1870 – December 6, 1952) was an American experimental

biologist A biologist is a scientist who conducts research in biology. Biologists are interested in studying life on Earth, whether it is an individual Cell (biology), cell, a multicellular organism, or a Community (ecology), community of Biological inter ...

who made seminal contributions to the genetics of the fruit fly ''

Drosophila melanogaster ''Drosophila melanogaster'' is a species of fly (the taxonomic order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly" or "pomace fly". Starting with Ch ...

'', although her work was obscured by the attention given her husband, Nobel laureate

Thomas Hunt Morgan Thomas Hunt Morgan (September 25, 1866 – December 4, 1945) was an American evolutionary biologist, geneticist, embryologist, and science author who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries elucidating the role tha ...

. Lilian Morgan published sixteen single-author papers between 1894 and 1947. Probably her most significant scientific contribution was the discovery of the attached-X chromosome and an entirely new pattern of inheritance in ''Drosophila'' in 1921. She also discovered the closed or ring-X chromosome in 1933. Both are important research tools today.

Early life

Morgan was born in 1870 in

Hallowell, Maine Hallowell is a city in Kennebec County, Maine, United States. The population was 2,570 at the 2020 census. Popular with tourists, Hallowell is noted for its culture and old architecture. Hallowell is included in the Augusta, Maine, micropolita ...

. She was orphaned at the age of three when her parents and younger sister died of

tuberculosis Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, in ...

. After the death of her parents, Morgan and her older sister Edith were raised by her maternal grandparents in Germantown, Pennsylvania.

Early research career

Morgan enrolled as an undergraduate student at Bryn Mawr in 1887. She majored in biology and was advised by

Martha Carey Thomas Martha Carey Thomas (January 2, 1857 – December 2, 1935) was an American educator, suffragist, and linguist. She was the second president of Bryn Mawr College, a women's liberal arts college in Bryn Mawr, Pennsylvania. Biography Early life ...

. After graduating with honors in 1891, she spent the summer at the

Marine Biological Laboratory The Marine Biological Laboratory (MBL) is an international center for research and education in biological and environmental science. Founded in Woods Hole, Massachusetts, in 1888, the MBL is a private, nonprofit institution that was independent ...

Woods Hole Woods Hole is a census-designated place in the town of Falmouth in Barnstable County, Massachusetts, United States. It lies at the extreme southwest corner of Cape Cod, near Martha's Vineyard and the Elizabeth Islands. The population was 781 at ...

, Massachusetts, where

Edmund Beecher Wilson Edmund Beecher Wilson (October 19, 1856 – March 3, 1939) was a pioneering American zoologist and geneticist. He wrote one of the most influential textbooks in modern biology, ''The Cell''. Career Wilson was born in Geneva, Illinois, the so ...

, one of her previous

zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...

professors, introduced her to her future graduate advisor and husband, Thomas Hunt Morgan. In the autumn of 1891, a Morgan earned a fellowship, which enabled her to study the musculature of

chiton Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...

s at the

University of Zurich The University of Zürich (UZH, german: Universität Zürich) is a public research university located in the city of Zürich, Switzerland. It is the largest university in Switzerland, with its 28,000 enrolled students. It was founded in 1833 f ...

with

Arnold Lang Arnold Lang (18 June 1855 – 30 November 1914) was a Swiss naturalist, a comparative anatomist and student of German biologist Ernst Haeckel. Biography In March 1876 he earned his PhD from the University of Jena, and two months later became hab ...

, a

comparative anatomist Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in t ...

and student of

Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...

. She returned to Bryn Mawr in 1892, where she earned a Master of Science in biology in 1894, under the advisory of Thomas Morgan. After graduation, she published her work on the musculature of chitons, returned to Woods Hole as an independent investigator, and spent many summers investigating breeding, development and

embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, '' -logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos and ...

amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...

Family life

In 1904, at the age of 34, she married Thomas Hunt Morgan and moved to

New York City New York, often called New York City or NYC, is the List of United States cities by population, most populous city in the United States. With a 2020 population of 8,804,190 distributed over , New York City is also the L ...

, where Thomas Morgan began working at

Columbia University Columbia University (also known as Columbia, and officially as Columbia University in the City of New York) is a private research university in New York City. Established in 1754 as King's College on the grounds of Trinity Church in Manhatt ...

. They spent the following summer in California, where she researched and published work on

planarian A planarian is one of the many flatworms of the traditional class Turbellaria. It usually describes free-living flatworms of the order Tricladida (triclads), although this common name is also used for a wide number of free-living platyhelmint ...

regeneration at the Stanford Marine Laboratory. She would not publish another paper for sixteen years. During the years following her marriage, she supported her husband's career and raised four children: Howard Key Morgan, born 1906; Edith Sampson Morgan, born 1907; Lilian Vaughan Morgan, born 1910; and Isabella Merrick Morgan, born 1911. Shine and Wrobel (1976) note that one key to

's success was that his personal affairs were entirely handled by Lilian Morgan, freeing him to focus on his research. The family spent their winters in New York and returned in the summers to Woods Hole, where she maintained a summer house for her children, relatives and her husband's graduate students. She maintained this house for many years, eventually equipping it for science lessons for children.

Involvement in science education

With several other women, Morgan founded the Summer School Club at Woods Hole in 1913, which is now the Children's School of Science, and served as its first educational chairperson and Science Committee Chair in 1914. She preferred working outdoors with children to conduct experiments and discuss problems.

Later research career

After her children were old enough, Morgan returned to the laboratory to study ''Drosophila'' genetics. Her husband, T.H. Morgan, encouraged her but did not collaborate with her. Instead, he gave her working space in his laboratory, called the "Fly Room," at

, where she maintained her own ''Drosophila'' stocks but held no official position. Her husband and the other male scientists never became comfortable with her presence in the lab, whose atmosphere was "a little like that of an exclusive men's club." Morgan may also have felt isolated because she was older than the other women and was neither outgoing nor talkative, according to

Alfred Sturtevant Alfred Henry Sturtevant (November 21, 1891 – April 5, 1970) was an American geneticist. Sturtevant constructed the first genetic map of a chromosome in 1911. Throughout his career he worked on the organism ''Drosophila melanogaster'' with ...

. Because she didn't hold an official position, she never attended a scientific meeting and never presented a paper at a conference.

Major research accomplishments

The attached-X chromosome

While working in the Fly Room at Columbia University, Morgan spotted an unusual fruit fly. She captured it and mated it with a normal male. She analyzed her data and discovered both an unusual chromosome and a new pattern of inheritance. This discovery became a powerful tool for X chromosome analysis. It provided further confirmation of the chromosome theory, sex determination, the linkage of traits on sex chromosomes, and an important tool for isolating and preserving traits on the X chromosome. After 100 years the attached-X strain continues to be used in genetics research and the new inheritance pattern, non-criss-cross inheritance, is taught in genetics courses and illustrated in genetics textbooks.

The ring-X chromosome

Another major contribution to ''Drosophila'' genetics was Lilian Morgan's discovery of

ring chromosomes A ring chromosome is an aberrant chromosome whose ends have fused together to form a ring. Ring chromosomes were first discovered by Lilian Vaughan Morgan in 1926. A ring chromosome is denoted by the symbol ''r'' in human genetics and ''R'' in ''D ...

. Ring chromosomes were discovered from their unusual frequencies of recombination in an attached-X stock.

Cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...

studies revealed a circularized X-chromosome. Ring-X chromosomes are unstable in early development and result in genetic mosaics, individuals with cells with differing genetic makeups. In females, for example, some cells could be XO and others XX. Ring chromosomes are a very important tool in the study of development.

Later life

Morgan and her family moved to California in 1928, where she continued her ''Drosophila'' research at the

California Institute of Technology The California Institute of Technology (branded as Caltech or CIT)The university itself only spells its short form as "Caltech"; the institution considers other spellings such a"Cal Tech" and "CalTech" incorrect. The institute is also occasional ...

in Pasadena while her husband

became the division head. Her husband died in 1945; one year afterwards, Morgan received her first official appointment as a research associate at the age of 76. She died in 1952 at the age of 82 in

Los Angeles Los Angeles ( ; es, Los Ángeles, link=no , ), often referred to by its initials L.A., is the largest city in the state of California and the second most populous city in the United States after New York City, as well as one of the world' ...

Publications

*Sampson, L. V. 1894. Die Muskulatur von Chiton. Jenaischen Zeitschrift für Naturwissenschaft 28: 460–468. *Sampson, L. V. 1895. The musculature of chiton. Journal of Morphology 11:595-628. *Sampson, L. V. 1900. Unusual modes of breeding and development among anura. Amer. Naturalist 34:687-715. *Sampson, L. V. 1904. A contribution to the embryology of Hylodes martinicensis. Araer. J. Anat. 3: 473–504. *Morgan, L. V. 1905. Incomplete anterior regeneration in the absence of the brain in Leploplana litloralis. Biol. Bull. 9:187-193. *Morgan, L. V. 1906. Regeneration of grafted pieces of planarians. J. Exp. Zool. 3:269-294. *Morgan, L. V. 1922. Non-criss-cross inheritance in Drosophila melanogaster. Biol. Bull. 42:267-274. *Morgan, L. V. 1925. Polyploidy in Drosophila melanogaster with two attached X chromosomes" ''Genetics'' 10:148-178. *Morgan, L. V. 1926. Correlation between shape and behavior of a chromosome" ''Proc. Natl. Acad. Sci'' 12:180-181. *Morgan, L. V. 1929. Composites of Drosophila melanogaster. Carnegie Inst. of Wash. Publ. No. 399: 225–296. *Morgan, L. V. 1931. Proof that bar changes to notbar by unequal crossing-over" ''Proc. Natl. Acad. Sci'' 17:270-272. *Morgan, L. V. 1933. A closed X chromosome in Drosophila melanogaster" ''Genetics'' 18:250-283. *Morgan, L. V. 1938a. Origin of attached-X chromosomes in Drosophila melanogaster and the occurrence of non-disjunction of X's in the male. Amer. Naturalist 72:434-446. *Morgan, L. V. 1938b. Effects of a compound duplication of the X chromosome of Drosophila melanogaster" ''Genetics'' 23:423-462. *Morgan, L. V. 1939. A spontaneous somatic exchange between non-homologous chromosomes in Drosophila melanogaster" ''Genetics'' 24:747-752. *Morgan, L. V. 1947. A variable phenotype associated with the fourth chromosome of Drosophila melanogaster and affected by heterochromatin" ''Genetics'' 32:200-219. *Morgan, T. H., H. Redfield, and L. V. Morgan. 1943. Maintenance of a Drosophila stock center, in connection with investigations on the germinal material in relation to heredity. Carnegie Inst. Wash. Yearbk. 42:171-174. *Morgan, T. H., A. H. Sturtevant, and L. V. Morgan. 1945. Maintenance of a Drosophila stock center, in connection with investigations on the germinal material in relation to heredity. Carnegie Inst. Wash. Yearbk. 44:157-160.

References

External links

Children's School of Science in Woods Hole, MA.
{{DEFAULTSORT:Morgan, Lilian Vaughn 1870 births 1952 deaths Morgan, Lilian Vaughan Bryn Mawr College alumni Bryn Mawr College faculty California Institute of Technology faculty Columbia University faculty People from Hallowell, Maine Scientists from New York City American women geneticists 19th-century American women scientists 20th-century American women scientists American women academics