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Thomas Hunt Morgan (September 25, 1866 – December 4, 1945) was an American embryologist who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries relating the role the chromosome plays in heredity. Morgan
received his PhD 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's research moved to the study of mutation in the fruit fly Drosophila
melanogaster. In his famous Fly Room at Columbia
University Morgan
was able to demonstrate that genes are carried on chromosomes and are the mechanical
basis of heredity. These discoveries formed the basis of the modern
science of genetics.
He was the first person to be awarded the Nobel Prize in Physiology or
Medicine for his work in genetics.
During his distinguished career, Morgan wrote 22
books and 370 scientific papers, and, as a result of his work, Drosophila became a major model
organism in contemporary genetics. The Division of
Biology he established at the California
Institute of Technology produced
seven Nobel Prize winners. Morgan
was born in Lexington, Kentucky,
to Charlton Hunt Morgan and Ellen Key Howard Morgan. Part of a long line of Southern aristocracy on his father's
side, Morgan was a nephew of Confederate General John Hunt Morgan and his great grandfather John Wesley Hunt had been the first
millionaire west of the Allegheny
Mountains. Through his mother, he was the great grandson of Francis Scott
Key, the author of the "Star Spangled
Banner", and John Eager
Howard, governor and senator from Maryland. However,
following the Civil War the family had fallen on harder times with the
loss of civil and property rights for those who aided the Confederacy.
His father also had difficulty finding work in politics and spent much
of his time coordinating veterans reunions. Beginning
at age 16 in the Preparatory Department, Morgan attended the State
College of Kentucky (now the University of
Kentucky). There, he focused on science; he particularly enjoyed
natural history, and worked with the U.S. Geological
Survey in his
summers. He graduated as valedictorian in 1886 with a bachelor in
science. Following a summer at the
Marine Biology School in Annisquam,
Massachusetts, Morgan began graduate studies in zoology at the recently founded Johns Hopkins
University, the first research oriented American university.
After two years of experimental work with morphologist William Keith
Brooks and
several publications, Morgan was eligible to receive a master of
science from the State College of Kentucky in 1888, the College
required two years 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 with respect to
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. With the fellowship he was
able to travel to Jamaica, the Bahamas and to Europe to conduct further research. In
1891, Morgan was appointed associate professor (and head of the biology
department) at Johns Hopkins' sister school Bryn Mawr
College, replacing his colleague E.B. Wilson.
Morgan taught all morphology related courses, while the other member of
the department, Jacques Loeb,
taught the physiological courses; though Loeb only stayed for one year,
it was the beginning of a lifelong friendship. Morgan
lectured in biology 5 days a week, giving two lectures a day. He
frequently included his own recent research in his lectures, and
although he was an enthusiastic teacher, his true interests were 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. At the laboratory in Naples
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. From his
exposure in Naples and through Loeb to the Entwicklungsmechanik (roughly, "developmental
mechanics") school of experimental biology — a reaction to the
vitalistic
Naturphilosophie that
was exteremly influential in 19th century morphology — Morgan's work
shifted from traditional, largely descriptive morphology to an
experimental embryology that sought physical and chemical explanations
for organismal development. 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 then 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. 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, work which was continued by Jacques Loeb (who became well known for
creating fatherless frogs using the method). Morgan
returned to Bryn Mawr in 1895 and 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,
published in 1897. He began a series of studies on different organisms
ability to regenerate. He looked at grafting and regeneration in
tadpoles, fish and earthworms and in 1901 this work was published as Regeneration.
Beginning in 1900, he started working on the problem of sex
determination; he also continued to study the evolutionary
problems that had been the focus of his earliest work. On June 4, 1904, Morgan married Lilian
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 in the
early years of their marriage, but would later contribute significantly
to Morgan's Drosophila work. Later in 1904, E.B. Wilson — still
blazing the path for his younger friend — invited Morgan to join him at Columbia
University, which at last
freed him to focus fully on experimental work. By
1904, when Morgan took a professorship in experimental zoology at Columbia
University, he was becoming increasingly focused on the
mechanisms of heredity and evolution. The previous year, he had
published Evolution
and Adaptation; like many biologists at that time, he saw clear
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 natural variation had distinct limits and
did not represent heritable changes. Embryological development posed an
additional problem in Morgan's view, as selection could not act on the
early, incomplete stages of highly complex organs such as the eye. The
common solution of the Lamarckian mechanism of inheritance of
acquired characters,
which featured prominently in Darwin's theory, was increasingly
rejected by biologists. According to Morgan's biographer Garland Allen,
he was also hindered by his views on taxonomy: he 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 he would remain skeptical of natural selection
for many years, his theories of heredity and variation were radically
transformed through his conversion to Mendelism. 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 had gone on to propose that new species are created by
mutation, bypassing the need for either Lamarckism or Darwinism. Morgan
dismissed both of these evolutionary theories, and was actually seeking
to prove Hugo
De Vries' mutation
theory with
his experimental heredity work. He was initially quite 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, and began cross-breeding
experiments to find heritable mutations. However, they had no
significant success for two years. Castle
had also had difficulty identifying mutations in Drosophila,
hardly unusual given the flies' tiny size. 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 bred with a red eyed female, their progeny
were all red eyed, while 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 the Belgian Professor Frans Alfons
Janssens of the University of
Leuven who
described the phenomenon in 1909 and had called it 'chiasmatypie'.
Morgan proposed that the amount of crossing over between linked genes
differs and that crossover frequency might indicate the distance
separating genes on the chromosome; later English geneticist J.B.S. Haldane suggested that the unit of
measurement for linkage be called the
morgan.
Morgan's student Alfred
Sturtevant developed the first genetic map in 1913. In
1915 Morgan, Sturtevant, Calvin Bridges and H.J. Muller wrote the seminal book The Mechanism of
Mendelian Heredity.
Geneticist Curt Stern called the book "the
fundamental textbook of the new genetics" and C.H. Waddington noted that "Morgan's theory
of the chromosome represents a great leap of imagination comparable
with Galileo or Newton". In the following years, most
biologists came to accept the Mendelian
chromosome theory, which was independently proposed by Walter Sutton and Theodor Boveri in 1902/1903 and elaborated
and expanded by Morgan and his students. Garland Allen characterized
the post 1915 period as one of normal science,
in which "The activities of 'geneticists' were aimed at further
elucidation of the details and implications of the Mendelian chromosome
theory developed between 1910 and 1915." However, the details of the
increasingly complex theory, as well as the very concept of the gene and its physical nature,
were still controversial. Critics such as W.E. Castle pointed to contrary results
in other organisms suggesting that genes interact with each other,
while to Richard
Goldschmidt and others, there was no compelling reason to view
genes as discrete units residing on chromosomes. 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. Morgan's group remained highly
productive, but Morgan largely withdrew from
doing fly work himself and gave his lab members considerable freedom in
designing and carrying out their own experiments. Instead, Morgan
returned to embryology and worked to encourage the spread of genetics
research to other organisms and the spread of the mechanistic
experimental approach (Enwicklungsmechanik) to all biological
fields. After 1915, he also became a
strong critic of the growing eugenics movement, which frequently
co-opted the ideas of genetics in support of racism and worse. John
Hopkins awarded Morgan an honorary LL.D. and the University of Kentucky
awarded him an honorary Ph.D. He was elected a member of the National
Academy of Sciences and made a foreign member of the Royal
Society. In 1924 Morgan
received the
Darwin Medal. His 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. 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,
and Sidney W. Fox. In
accordance with his reputation, Morgan held 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
acknowledgement of the group nature of his discovery he gave his prize
money to Bridges', Sturtevant's 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
counselling. 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 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. Morgan had throughout his life suffered
with a chronic duodenal
ulcer, and in 1945 he
experienced a severe heart attack and died from a ruptured artery. Morgan
was interested in evolution throughout his life. He wrote his thesis on
the phylogeny of sea spiders (pycnogonids)
and wrote 4 books about evolution. In Evolution
and Adaptation (1903)
he argued the anti-Darwinist position that selection never could
produce wholly new species by acting on slight individual differences. He also rejected Darwin's
theory of sexual selection and
the Neo-Lamarckian theory of the inheritance of acquired characters.
Morgan was not the only scientist attacking natural selection. The
period 1875 - 1925 has been called 'The eclipse of Darwinism'.
After discovering many small stable heritable mutations in Drosophila he
gradually changed his mind. The relevance of mutations for evolution is
that only characters that are inherited can have an effect in
evolution. Since Morgan (1915) 'solved the problem of heredity', he was
in a unique position to examine critically Darwin's theory of natural
selection. In A
Critique of the Theory of Evolution (1916) he 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 that "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". Injurious mutations have practically no
chance of becoming established.
Far from rejecting evolution as the title of his 1916 book may suggest,
Morgan not only laid the foundation of the science of genetics, but by
doing that, 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. Therefore, Darwinism could
not progress without a correct theory of genetics. Morgan created that
foundation and that is why Morgan's work is so fundamental for the neo-Darwinian synthesis
despite his criticism at the beginning of his career. Even so, much
work on the Evolutionary Synthesis remained to be done. 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." The
Thomas Hunt Morgan School of Biological Sciences at the University of
Kentucky is named for Morgan. In Morgan's honor, the Genetics
Society of America annually
awards the Thomas Hunt
Morgan Medal 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. |