Thomas Hunt Morgan

Thomas Hunt Morgan (September 25, 1866 –

December 4, 1945)[2] was an American evolutionary Thomas Hunt Morgan
ForMemRS
biologist, geneticist, embryologist, and science author
who won the Nobel Prize in Physiology or Medicine in
1933 for discoveries elucidating the role that the
chromosome plays in heredity.[3]

Morgan received his Ph.D. from Johns Hopkins

University in zoology in 1890 and researched
embryology during his tenure at Bryn Mawr.
Following the rediscovery of Mendelian inheritance in
1900, Morgan began to study the genetic
characteristics of the fruit fly Drosophila
melanogaster. In his famous Fly Room at Columbia
University's Schermerhorn Hall, Morgan demonstrated
that genes are carried on chromosomes and are the
mechanical basis of heredity. These discoveries formed
the basis of the modern science of genetics. Morgan in 1891
Born September 25, 1866
During his distinguished career, Morgan wrote 22 Lexington, Kentucky, US
books and 370 scientific papers.[2] As a result of his
Died December 4, 1945 (aged 79)
work, Drosophila became a major model organism in
Pasadena, California, US
contemporary genetics. The Division of Biology which
he established at the California Institute of Technology Alma mater University of Kentucky (B.S.)
has produced seven Nobel Prize winners. Johns Hopkins University (Ph.D.)

Known for Establishing Drosophila

melanogaster as a major model
Early life and education organism in genetics

Morgan was born in Lexington, Kentucky, to Charlton Linked genes

Hunt Morgan and Ellen Key Howard Morgan.[3][4] Awards Member of the National
Part of a line of Southern plantation and slave owners Academy of Sciences (1909)[1]
on his father's side, Morgan was a nephew of Foreign Member of the Royal
Confederate General John Hunt Morgan; his great- Society (1919)[2]
grandfather John Wesley Hunt had been one of the first
Nobel Prize in Physiology or
millionaires west of the Allegheny Mountains.
Medicine (1933)
Through his mother, he was the great-grandson of
Copley Medal (1939)
Francis Scott Key, the author of the "Star Spangled
Banner", and John Eager Howard, governor and Scientific career
senator from Maryland.[4] Following the Civil War, the Fields Genetics
family fell on hard times with the temporary loss of
Embryology
civil and some property rights for those who aided the Institutions Bryn Mawr College
Confederacy. His father had difficulty finding work in Columbia University
politics and spent much of his time coordinating
California Institute of
veterans' reunions.
Technology
Beginning at age 16 in the Preparatory Department, Marine Biological Laboratory
Morgan attended the State College of Kentucky (now Doctoral Nettie Maria Stevens
the University of Kentucky). He focused on science; he students
John Howard Northrop
particularly enjoyed natural history, and worked with
Hermann Joseph Muller
the U.S. Geological Survey in his summers. He
graduated as valedictorian in 1886 with a Bachelor of Calvin Bridges
[5]
Science degree. Following a summer at the Marine Alfred Sturtevant
Biology School in Annisquam, Massachusetts, Morgan Chester Ittner Bliss
began graduate studies in zoology at the recently Tan Jiazhen
founded Johns Hopkins University. After two years of
Signature
experimental work with morphologist William Keith
Brooks and writing several publications, Morgan was
eligible to receive a Master of Science from the State
College of Kentucky in 1888. The college required two
years of study at another institution and an examination by the college faculty. The college offered
Morgan a full professorship; however, he chose to stay at Johns Hopkins and was awarded a relatively
large fellowship to help him fund his studies.

Under Brooks, Morgan completed his thesis work on the embryology of sea spiders—collected during the
summers of 1889 and 1890 at the Marine Biological Laboratory in Woods Hole, Massachusetts—to
determine their phylogenetic relationship with other arthropods. He concluded that concerning
embryology, they were more closely related to spiders than crustaceans. Based on the publication of this
work, Morgan was awarded his Ph.D. from Johns Hopkins in 1890 and was also awarded the Bruce
Fellowship in Research. He used the fellowship to travel to Jamaica, the Bahamas and Europe to conduct
further research.[6]

Every summer from 1910 to 1925, Morgan and his colleagues at the famous Fly Room at Columbia
University moved their research program to the Marine Biological Laboratory. Aside from being an
independent investigator at the MBL from 1890 to 1942, he became very involved in the governance of
the institution, including serving as an MBL trustee from 1897 to 1945.[7]

Career and research

Bryn Mawr
In 1890, Morgan was appointed associate professor (and head of the biology department) at Johns
Hopkins' sister school Bryn Mawr College, replacing his colleague Edmund Beecher Wilson.[8] Morgan
taught all morphology-related courses, while the other member of the department, Jacques Loeb, taught
the physiological courses. Although Loeb stayed for only one year, it was the beginning of their lifelong
friendship.[9] Morgan lectured in biology five days a week, giving two lectures a day. He frequently
included his recent research in his lectures. Although an enthusiastic teacher, he was most interested in
research in the laboratory. During the first few years at Bryn Mawr, he produced descriptive studies of sea
acorns, ascidian worms, and frogs.

In 1894 Morgan was granted a year's absence to conduct research in the laboratories of Stazione
Zoologica in Naples, where Wilson had worked two years earlier. There he worked with German
biologist Hans Driesch, whose research in the experimental study of development piqued Morgan's
interest. Among other projects that year, Morgan completed an experimental study of ctenophore
embryology. In Naples and through Loeb, he became familiar with the Entwicklungsmechanik (roughly,
"developmental mechanics") school of experimental biology. It was a reaction to the vitalistic
Naturphilosophie, which was extremely influential in 19th-century morphology. Morgan changed his
work from traditional, largely descriptive morphology to experimental embryology that sought physical
and chemical explanations for organismal development.[10]

At the time, there was considerable scientific debate over the question of how an embryo developed.
Following Wilhelm Roux's mosaic theory of development, some believed that hereditary material was
divided among embryonic cells, which were predestined to form particular parts of a mature organism.
Driesch and others thought that development was due to epigenetic factors, where interactions between
the protoplasm and the nucleus of the egg and the environment could affect development. Morgan was in
the latter camp; his work with Driesch demonstrated that blastomeres isolated from sea urchin and
ctenophore eggs could develop into complete larvae, contrary to the predictions (and experimental
evidence) of Roux's supporters.[11] A related debate involved the role of epigenetic and environmental
factors in development; on this front Morgan showed that sea urchin eggs could be induced to divide
without fertilization by adding magnesium chloride. Loeb continued this work and became well known
for creating fatherless frogs using the method.[12] [13]

When Morgan returned to Bryn Mawr in 1895, he was promoted to full professor. Morgan's main lines of
experimental work involved regeneration and larval development; in each case, his goal was to
distinguish internal and external causes to shed light on the Roux-Driesch debate. He wrote his first book,
The Development of the Frog's Egg (1897). He began a series of studies on different organisms' ability to
regenerate. He looked at grafting and regeneration in tadpoles, fish, and earthworms; in 1901 he
published his research as Regeneration.

Beginning in 1900, Morgan started working on the problem of sex determination, which he had
previously dismissed when Nettie Stevens discovered the impact of the Y chromosome on sex. He also
continued to study the evolutionary problems that had been the focus of his earliest work.[14]

Columbia University
Morgan worked at Columbia University for 24 years, from 1904 until 1928 when he left for a position at
the California Institute of Technology.

In 1904, his friend, Jofi Joseph died of tuberculosis, and he felt he ought to mourn her, though E. B.
Wilson—still blazing the path for his younger friend—invited Morgan to join him at Columbia
University. This move freed him to focus fully on experimental work.[15]
When Morgan took the professorship in experimental zoology, he
became increasingly focused on the mechanisms of heredity and
evolution. He published Evolution and Adaptation (1903); like many
biologists at the time, he saw evidence for biological evolution (as in
the common descent of similar species) but rejected Darwin's
proposed mechanism of natural selection acting on small, constantly
produced variations.

Extensive work in biometry seemed to indicate that continuous In a typical Drosophila genetics
natural variation had distinct limits and did not represent heritable experiment, male and female
flies with known phenotypes are
changes. Embryological development posed an additional problem in
put in a jar to mate; females
Morgan's view, as selection could not act on the early, incomplete
must be virgins. Eggs are laid in
stages of highly complex organs such as the eye. The common porridge which the larvae feed
solution of the Lamarckian mechanism of inheritance of acquired on; when the life cycle is
characters, which featured prominently in Darwin's theory, was complete, the progeny are
increasingly rejected by biologists. According to Morgan's biographer scored for the inheritance of the
Garland Allen, he was also hindered by his views on taxonomy: he trait of interest.
thought that species were entirely artificial creations that distorted the
continuously variable range of real forms, while he held a
"typological" view of larger taxa and could see no way that one such group could transform into another.
But while Morgan was skeptical of natural selection for many years, his theories of heredity and variation
were radically transformed through his conversion to Mendelism.[16]

In 1900 three scientists, Carl Correns, Erich von Tschermak and Hugo De Vries, had rediscovered the
work of Gregor Mendel, and with it the foundation of genetics. De Vries proposed that new species were
created by mutation, bypassing the need for either Lamarckism or Darwinism. As Morgan had dismissed
both evolutionary theories, he was seeking to prove De Vries' mutation theory with his experimental
heredity work. He was initially skeptical of Mendel's laws of heredity (as well as the related
chromosomal theory of sex determination), which were being considered as a possible basis for natural
selection.

Following C. W. Woodworth and William E. Castle, around

1908 Morgan started working on the fruit fly Drosophila
melanogaster, and encouraging students to do so as well.
With Fernandus Payne, he mutated Drosophila through
physical, chemical, and radiational means.[17][18] He began
cross-breeding experiments to find heritable mutations, but
they had no significant success for two years.[17] Castle had
also had difficulty identifying mutations in Drosophila, which
were tiny. Finally, in 1909, a series of heritable mutants
appeared, some of which displayed Mendelian inheritance
patterns; in 1910 Morgan noticed a white-eyed mutant male
among the red-eyed wild types. When white-eyed flies were Sex linked inheritance of the white eyed
bred with a red-eyed female, their progeny were all red-eyed. mutation.
A second-generation cross produced white-eyed males—a
sex-linked recessive trait, the gene for which Morgan named
white. Morgan also discovered a pink-eyed mutant that showed a different pattern of inheritance. In a
paper published in Science in 1911, he concluded that (1) some traits were sex-linked, (2) the trait was
probably carried on one of the sex chromosomes, and (3) other genes were probably carried on specific
chromosomes as well.

Morgan and his students became more successful at finding

mutant flies; they counted the mutant characteristics of thousands
of fruit flies and studied their inheritance. As they accumulated
multiple mutants, they combined them to study more complex
inheritance patterns. The observation of a miniature-wing mutant,
which was also on the sex chromosome but sometimes sorted
independently to the white-eye mutation, led Morgan to the idea of
genetic linkage and to hypothesize the phenomenon of crossing
over. He relied on the discovery of Frans Alfons Janssens, a
Morgan's illustration of crossing
Belgian professor at the University of Leuven, who described the
over, from his 1916 A Critique of the
phenomenon in 1909 and had called it chiasmatypy. Morgan Theory of Evolution
proposed that the amount of crossing over between linked genes
differs and that crossover frequency might indicate the distance
separating genes on the chromosome. The later English geneticist
J. B. S. Haldane suggested that the unit of measurement for First genetic map (Sturtevant,
linkage be called the morgan. Morgan's student Alfred Sturtevant 1913). It shows 6 sex-linked genes.
developed the first genetic map in 1913.[19]

In 1915 Morgan, Sturtevant, Calvin Bridges and H. J. Muller

wrote the seminal book The Mechanism of Mendelian
Heredity.[21] Geneticist Curt Stern called the book "the
fundamental textbook of the new genetics".[22]

In the following years, most biologists came to accept the

Mendelian-chromosome theory, which was independently Thomas Hunt Morgan's Drosophila
proposed by Walter Sutton and Theodor Boveri in 1902/1903, melanogaster genetic linkage map. This
and elaborated and expanded by Morgan and his students. was the first successful gene mapping
work and provides important evidence for
Garland Allen characterized the post-1915 period as one of
the chromosome theory of inheritance.
normal science, in which "The activities of 'geneticists' were The map shows the relative positions of
aimed at further elucidation of the details and implications of allelic characteristics on the second
the Mendelian-chromosome theory developed between 1910 Drosophila chromosome. The distance
and 1915." But, the details of the increasingly complex between the genes (map units) is equal
theory, as well as the concept of the gene and its physical to the percentage of crossing-over events
nature, were still controversial. Critics such as W. E. Castle that occurs between different alleles.[20]
pointed to contrary results in other organisms, suggesting that
genes interact with each other, while Richard Goldschmidt
and others thought there was no compelling reason to view genes as discrete units residing on
chromosomes.[24]

Because of Morgan's dramatic success with Drosophila, many other labs throughout the world took up
fruit fly genetics. Columbia became the center of an informal exchange network, through which
promising mutant Drosophila strains were transferred from lab to lab; Drosophila became one of the first
and for some time the most widely used, model organisms.[25]
Morgan's group remained highly productive, but Morgan largely
withdrew from doing fly work and gave his lab members
considerable freedom in designing and carrying out their own
experiments.

He returned to embryology and worked to encourage the spread of

genetics research to other organisms and the spread of mechanistic
experimental approach (Enwicklungsmechanik) to all biological
fields.[26] After 1915, he also became a strong critic of the
growing eugenics movement, which adopted genetic approaches
in support of racist views of "improving" humanity.[27]

Morgan's fly-room at Columbia became world-famous, and he

found it easy to attract funding and visiting academics. In 1927
after 25 years at Columbia, and nearing the age of retirement, he
received an offer from George Ellery Hale to establish a school of
biology in California.

Caltech Genetic map of drosophila,

published in The theory of the gene
In 1928 Morgan joined the faculty of the California Institute of 1926 edition.[23]
Technology where he remained until his retirement 14 years later
in 1942.

Morgan moved to California to head the Division of Biology at the

California Institute of Technology in 1928. In establishing the
biology division, Morgan wanted to distinguish his program from
those offered by Johns Hopkins and Columbia, with research
focused on genetics and evolution; experimental embryology;
physiology; biophysics, and biochemistry. He was also
instrumental in the establishment of the Marine Laboratory at
Corona del Mar. He wanted to attract the best people to the
Division at Caltech, so he took Bridges, Sturtevant, Jack Shultz
and Albert Tyler from Columbia and took on Theodosius
Dobzhansky as an international research fellow. More scientists
came to work in the Division including George Beadle, Boris
Ephrussi, Edward L. Tatum, Linus Pauling, Frits Went, Edward B.
1931 drawing of Thomas Hunt
Lewis, and Sidney W. Byance with his reputation, Morgan held
Morgan
numerous prestigious positions in American science organizations.
From 1927 to 1931 Morgan served as the President of the National
Academy of Sciences; in 1930 he was the President of the American Association for the Advancement of
Science; and in 1932 he chaired the Sixth International Congress of Genetics in Ithaca, New York. In
1933 Morgan was awarded the Nobel Prize in Physiology or Medicine; he had been nominated in 1919
and 1930 for the same work. As an acknowledgment of the group nature of his discovery, he gave his
prize money to Bridges, Sturtevant, and his own children. Morgan declined to attend the awards
ceremony in 1933, instead attending in 1934. The 1933 rediscovery of the giant polytene chromosomes in
the salivary gland of Drosophila may have influenced his choice. Until that point, the lab's results had
been inferred from phenotypic results, the visible polytene chromosome enabled them to confirm their
results on a physical basis. Morgan's Nobel acceptance speech entitled "The Contribution of Genetics to
Physiology and Medicine" downplayed the contribution genetics could make to medicine beyond genetic
counseling. In 1939 he was awarded the Copley Medal by the Royal Society.

He received two extensions of his contract at Caltech, but eventually retired in 1942, becoming a
professor and chairman emeritus. George Beadle returned to Caltech to replace Morgan as chairman of
the department in 1946. Although he had retired, Morgan kept offices across the road from the Division
and continued laboratory work. In his retirement, he returned to the questions of sexual differentiation,
regeneration, and embryology.

Death
Morgan had throughout his life suffered from a chronic duodenal ulcer. In 1945, at age 79, he
experienced a severe heart attack and died from a ruptured artery.

Morgan and evolution

Morgan was interested in evolution throughout his life. He wrote his thesis on the phylogeny of sea
spiders (pycnogonids) and wrote four books about evolution. In Evolution and Adaptation (1903), he
argued the anti-Darwinist position that selection could never produce wholly new species by acting on
slight individual differences.[28] He rejected Darwin's theory of sexual selection[29] and the Neo-
Lamarckian theory of the inheritance of acquired characters.[30] Morgan was not the only scientist
attacking natural selection. The period 1875–1925 has been called 'The eclipse of Darwinism'.[31] After
discovering many small stable heritable mutations in Drosophila, Morgan gradually changed his mind.
The relevance of mutations for evolution is that only characters that are inherited can have an effect on
evolution. Since Morgan solved the problem of heredity (1915), he was in a unique position to examine
critically Darwin's theory of natural selection.

In A Critique of the Theory of Evolution (1916), Morgan discussed questions such as: "Does selection
play any role in evolution? How can selection produce anything new? Is selection no more than the
elimination of the unfit? Is selection a creative force?" After eliminating some misunderstandings and
explaining in detail the new science of Mendelian heredity and its chromosomal basis, Morgan concludes,
"the evidence shows clearly that the characters of wild animals and plants, as well as those of
domesticated races, are inherited both in the wild and in domesticated forms according to the Mendel's
Law". "Evolution has taken place by the incorporation into the race of those mutations that are beneficial
to the life and reproduction of the organism".[32] Injurious mutations have practically no chance of
becoming established.[33] Far from rejecting evolution, as the title of his 1916 book may suggest,
Morgan, laid the foundation of the science of genetics. He also laid the theoretical foundation for the
mechanism of evolution: natural selection. Heredity was a central plank of Darwin's theory of natural
selection, but Darwin could not provide a working theory of heredity. Darwinism could not progress
without a correct theory of genetics. By creating that foundation, Morgan contributed to the neo-
Darwinian synthesis, despite his criticism of Darwin at the beginning of his career. Much work on the
Evolutionary Synthesis remained to be done.

Awards and honors

Morgan left an important legacy in genetics. Some of Morgan's students from Columbia and Caltech went
on to win their own Nobel Prizes, including George Wells Beadle and Hermann Joseph Muller. Nobel
prize winner Eric Kandel has written of Morgan, "Much as Darwin's insights into the evolution of animal
species first gave coherence to nineteenth-century biology as a descriptive science, Morgan's findings
about genes and their location on chromosomes helped transform biology into an experimental
science."[34]

Johns Hopkins awarded Morgan an honorary LL.D. and the University of Kentucky awarded
him an honorary Ph.D.
He was elected Member of the National Academy of Sciences in 1909.[1]
He was elected to the American Philosophical Society in 1915.[35]
He was elected a Foreign Member of the Royal Society (ForMemRS) in 1919[2]
In 1924 Morgan received the Darwin Medal.
He was elected to the American Academy of Arts and Sciences in 1928.[36]
The Thomas Hunt Morgan School of Biological Sciences at the University of Kentucky is
named for him.
The Genetics Society of America annually awards the Thomas Hunt Morgan Medal, named
in his honor, to one of its members who has made a significant contribution to the science of
genetics.
Thomas Hunt Morgan's discovery was illustrated on a 1989 stamp issued in Sweden,
showing the discoveries of eight Nobel Prize-winning geneticists.
A junior high school in Shoreline, Washington was named in Morgan's honor for the latter
half of the 20th century.

Personal life
On June 4, 1904, Morgan married Lillian Vaughan Sampson (1870–1952), who had entered graduate
school in biology at Bryn Mawr the same year Morgan joined the faculty; she put aside her scientific
work for 16 years of their marriage when they had four children. Later she contributed significantly to
Morgan's Drosophila work. One of their four children (one boy and three girls) was Isabel Morgan
(1911–1996) (Marr. Mountain), who became a virologist at Johns Hopkins, specializing in polio research.
Morgan was an atheist.[37][38][39][40]

See also
Mildred Hoge Richards, pupil

References
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5. Allen (1978), pp. 11–14, 24.
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ww.stanford.edu/group/Urchin/loeb.htm). American Journal of Physiology. 31 (3): 135–138.
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artificialparth00loebgoog). University of Chicago Press. "jacques loeb sea urchin."
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17. Kohler, Lords of the Fly, pp. 37–43
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25. Kohler, Lords of the Fly, chapter 5
26. Allen, Thomas Hunt Morgan, pp. 214–215, 285
27. Allen, Thomas Hunt Morgan, pp. 227–234
28. Allen, Garland E. (2009). Ruse, Michael; Travis, Joseph (eds.). Evolution. The First Four
Billion Years (https://archive.org/details/evolutionfirstfo00mich/page/746). Harvard University
Press. p. 746 (https://archive.org/details/evolutionfirstfo00mich/page/746).
ISBN 9780674031753.
29. "I think we shall be justified in rejecting it as an explanation of the secondary sexual
differences amongst animals", pp. 220–221, chapter VI, Evolution and Adaptation, 1903.
30. Chapter VII of Evolution and Adaptation, 1903.
31. Bowler, Peter (2003). Evolution. The History of an Idea. University of California Press.
chapter 7.
32. A Critique of the Theory of Evolution, Princeton University Press, 1916, pp. 193–194
33. A Critique of the Theory of Evolution, p. 189.
34. Kandel, Eric. 1999. "Genes, Chromosomes, and the Origins of Modern Biology" (http://www.
columbia.edu/cu/alumni/Magazine/Legacies/Morgan/), Columbia Magazine
35. "APS Member History" (https://search.amphilsoc.org/memhist/search?creator=Thomas+H.+
Morgan&title=&subject=&subdiv=&mem=&year=&year-max=&dead=&keyword=&smode=ad
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Academy of Arts & Sciences. 2023-02-09. Retrieved 2023-11-02.
37. George Pendle (2006). Strange Angel: The Otherworldly Life of Rocket Scientist John
Whiteside Parsons. Houghton Mifflin Harcourt. p. 69. ISBN 9780156031790. "The Nobel
Prize-winning geneticist and stringent atheist Thomas Hunt Morgan was developing the
chromosome theory of heredity by examining his swarm of mutated Drosophila (fruit flies)
through a jeweler's loupe."
38. "Morgan's passion for experimentation was symptomatic of his general skepticism and his
distaste for speculation. He believed only what could be proven. He was said to be an
atheist, and I have always believed that he was. Everything I knew about him—his
skepticism, his honesty—was consistent with disbelief in the supernatural." Norman H.
Horowitz, T. H. Morgan at Caltech: A Reminiscence (http://www.genetics.org/content/149/4/1
629.short), Genetics, Vol. 149, 1629–1632, August 1998.
39. Judith R. Goodstein. "The Thomas Hunt Morgan Era in Biology" (https://web.archive.org/we
b/20160822085324/http://calteches.library.caltech.edu/3677/1/Goodstein.pdf) (PDF).
Calteches.library.caltech.edu. Archived from the original (http://calteches.library.caltech.edu/
3677/1/Goodstein.pdf) (PDF) on 22 August 2016. Retrieved 28 April 2019.
40. Horowitz, Norman H. (1 August 1998). "T. H. Morgan at Caltech: A Reminiscence" (https://w
eb.archive.org/web/20160405111515/http://www.genetics.org/content/149/4/1629).
Genetics. 149 (4): 1629–1632. doi:10.1093/genetics/149.4.1629 (https://doi.org/10.1093%2
Fgenetics%2F149.4.1629). PMC 1460264 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1
460264). PMID 9691024 (https://pubmed.ncbi.nlm.nih.gov/9691024). Archived from the
original (http://www.genetics.org/content/149/4/1629) on 5 April 2016. Retrieved 6 February
2017.

Further reading
Allen, Garland E. (1978). Thomas Hunt Morgan: The Man and His Science. Princeton
 University Press. ISBN 0-691-08200-6.
Allen, Garland E. (2000). "Morgan, Thomas Hunt". American National Biography. Oxford
University Press.
Kohler, Robert E. (1994). Lords of the Fly: Drosophila Genetics and the Experimental Life (h
ttps://archive.org/details/lordsofflydrosop0000kohl). University of Chicago Press. ISBN 0-
226-45063-5.
Shine, Ian B; Sylvia Wrobel (1976). Thomas Hunt Morgan: Pioneer of Genetics. University
Press of Kentucky. ISBN 0-8131-0095-X.
Stephenson, Wendell H. (April 1946). "Thomas Hunt Morgan: Kentucky's Gift to Biological
Science" (http://connect1.ajaxdocumentviewer.com/viewerajax.php?YH6zlSpldS1Kk8DNWc
zPQfWRQEZaQiigWtcQqyAJPOzwCuENy3X9NEGHGfp5ezCHJdgdN49hce5zRYKQV9Ooz
8GIMU6APbD3Q44QsntYrbvmOfVZcl%2Fc%2F%2F5A%2F7UCCAINs5TDdf9GrHiUxgTSH
v2Jd52MmR0%2F6g%2BvmyEeqX%2BT749vkDd6hg8ex8I%2Blw72Drk7SZGOzSYgb9%2
F%2BUfLtOqzVmePrUKPKEE9t36zcvEkB%2BvXI9s5sm1%2F7h3zK02hRslDLJrcdjcGmmk
Y0fztmIHiQNPDYl83%2FUr4vLG8T8goUT7L5s97UVjU2ZXl6aCrbfElXaHhEzcVOZP1oTfgz
QVHJqtBxl%2BAOUqfEitnUEF1O3RyGXrogBGYt0fuLdiWLorXwxBK1T32%2FNtWmvGjLE
TcouG9zhPWncybvoRFWp%2FsX%2BQk%3D). Filson Club History Quarterly. 20 (2).
Retrieved 2012-02-22.
Sturtevant, Alfred H. (1959). "Thomas Hunt Morgan". Biographical Memoirs of the National
Academy of Sciences. 33: 283–325.

External links
Thomas Hunt Morgan (https://www.nobelprize.org/laureate/325) on Nobelprize.org including
the Nobel Lecture on June 4, 1934 The Relation of Genetics to Physiology and Medicine
Thomas Hunt Morgan Biological Sciences Building at University of Kentucky (http://ukcc.uk
y.edu/cgi-bin/dynamo?maps.391+campus+0225)
Thomas Hunt Morgan (https://web.archive.org/web/20090211065202/http://nobelmedicine.c
o.uk/thomashuntmorgan.htm)
Thomas Hunt Morgan (https://www.nasonline.org/wp-content/uploads/2024/06/morgan-thom
as-hunt.pdf) – Biographical Memoirs (https://www.nasonline.org/publications/biographical-m
emoirs/) of the National Academy of Sciences
Works by Thomas Hunt Morgan (https://www.gutenberg.org/ebooks/author/34763) at Project
Gutenberg
Works by or about Thomas Hunt Morgan (https://archive.org/search.php?query=%28%28su
bject%3A%22Morgan%2C%20Thomas%20Hunt%22%20OR%20subject%3A%22Morgan%
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2E%22%20OR%20subject%3A%22Thomas%20Hunt%20Morgan%22%20OR%20subject%
3A%22Thomas%20H%2E%20Morgan%22%20OR%20subject%3A%22T%2E%20H%2E%
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eator%3A%22Thomas%20H%2E%20Morgan%22%20OR%20creator%3A%22T%2E%20
H%2E%20Morgan%22%20OR%20creator%3A%22T%2E%20Hunt%20Morgan%22%20O
R%20creator%3A%22Morgan%2C%20Thomas%20Hunt%22%20OR%20creator%3A%22M
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H%2E%20Morgan%22%20OR%20title%3A%22T%2E%20H%2E%20Morgan%22%20OR%
20description%3A%22Thomas%20Hunt%20Morgan%22%20OR%20description%3A%22Th
omas%20H%2E%20Morgan%22%20OR%20description%3A%22T%2E%20H%2E%20Mor
gan%22%20OR%20description%3A%22Morgan%2C%20Thomas%20Hunt%22%20OR%2
0description%3A%22Morgan%2C%20Thomas%20H%2E%22%29%20OR%20%28%22186
 6-1945%22%20AND%20Morgan%29%29%20AND%20%28-mediatype:software%29) at
the Internet Archive
Works by Thomas Hunt Morgan (https://librivox.org/author/15789) at LibriVox (public domain
audiobooks)

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