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Thomas Hunt Morgan

 
Thomas Hunt Morgan

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Thomas Hunt Morgan



 
 
Thomas Hunt Morgan (September 25, 1866 – December 4, 1945) was an American geneticist
Genetics

Genetics , a discipline of biology, is the science of heredity and Genetic variation in living organisms. The fact that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding....
 and embryologist
Embryology

Embryology is the study of the development of an embryo. An embryo is defined as any organism in a stage before birth or hatching, or in plants, before germination occurs....
. Morgan received his PhD from Johns Hopkins University
Johns Hopkins University

The Johns Hopkins University, commonly referred to as Hopkins or JHU, is a private university research university located in Baltimore, Maryland, Maryland, United States....
 in 1890 and researched embryology during his tenure at Bryn Mawr
Bryn Mawr College

'Bryn Mawr College' is a highly selective Women's colleges in the United States Liberal arts colleges in the United States located in Bryn Mawr, Pennsylvania, a community in Lower Merion Township, Pennsylvania, Pennsylvania, ten miles west of Philadelphia, Pennsylvania....
. Following the rediscovery of Mendelian inheritance
Mendelian inheritance

Mendelian inheritance is a set of primary tenets relating to the transmission of heredity characteristics from parent organisms to their children; it underlies much of genetics....
 in 1900, Morgan's research moved to the study of mutation
Mutation

In biology, mutations are changes to the nucleotide sequence of the genetic material of an organism. Mutations can be caused by copying errors in the genetic material during cell division, by exposure to ultraviolet or ionizing radiation, chemical mutagens, or virus , or can be induced by the organism, itself, by cellular processes such as s...
 in the fruit fly Drosophila melanogaster
Drosophila melanogaster

Drosophila melanogaster is a two-winged insect that belongs to the Diptera, the Order of the Fly. The species is commonly known as the Drosophilidae or vinegar fly, and is one of the most commonly used model organisms in biology, including studies in genetics, physiology and Life history theory....
. In his famous Fly Room at Columbia University
Columbia University

Columbia University in the City of New York , is a private university in the United States and a member of the Ivy League. Columbia's main campus lies in the Morningside Heights, Manhattan neighborhood in the borough of Manhattan, in New York City....
 Morgan was able to demonstrate that gene
Gene

A gene is the basic unit of heredity in a living organism. All living things depend on genes. Genes hold the information to build and maintain their cell and pass genetic trait to offspring....
s are carried on chromosome
Chromosome

A chromosome is an organized structure of DNA and protein that is found in Cell . A chromosome is a single piece of DNA that contains many genes, regulatory sequence and other genetic sequence....
s and are the mechanical basis of heredity.






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Thomas Hunt Morgan (September 25, 1866 – December 4, 1945) was an American geneticist
Genetics

Genetics , a discipline of biology, is the science of heredity and Genetic variation in living organisms. The fact that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding....
 and embryologist
Embryology

Embryology is the study of the development of an embryo. An embryo is defined as any organism in a stage before birth or hatching, or in plants, before germination occurs....
. Morgan received his PhD from Johns Hopkins University
Johns Hopkins University

The Johns Hopkins University, commonly referred to as Hopkins or JHU, is a private university research university located in Baltimore, Maryland, Maryland, United States....
 in 1890 and researched embryology during his tenure at Bryn Mawr
Bryn Mawr College

'Bryn Mawr College' is a highly selective Women's colleges in the United States Liberal arts colleges in the United States located in Bryn Mawr, Pennsylvania, a community in Lower Merion Township, Pennsylvania, Pennsylvania, ten miles west of Philadelphia, Pennsylvania....
. Following the rediscovery of Mendelian inheritance
Mendelian inheritance

Mendelian inheritance is a set of primary tenets relating to the transmission of heredity characteristics from parent organisms to their children; it underlies much of genetics....
 in 1900, Morgan's research moved to the study of mutation
Mutation

In biology, mutations are changes to the nucleotide sequence of the genetic material of an organism. Mutations can be caused by copying errors in the genetic material during cell division, by exposure to ultraviolet or ionizing radiation, chemical mutagens, or virus , or can be induced by the organism, itself, by cellular processes such as s...
 in the fruit fly Drosophila melanogaster
Drosophila melanogaster

Drosophila melanogaster is a two-winged insect that belongs to the Diptera, the Order of the Fly. The species is commonly known as the Drosophilidae or vinegar fly, and is one of the most commonly used model organisms in biology, including studies in genetics, physiology and Life history theory....
. In his famous Fly Room at Columbia University
Columbia University

Columbia University in the City of New York , is a private university in the United States and a member of the Ivy League. Columbia's main campus lies in the Morningside Heights, Manhattan neighborhood in the borough of Manhattan, in New York City....
 Morgan was able to demonstrate that gene
Gene

A gene is the basic unit of heredity in a living organism. All living things depend on genes. Genes hold the information to build and maintain their cell and pass genetic trait to offspring....
s are carried on chromosome
Chromosome

A chromosome is an organized structure of DNA and protein that is found in Cell . A chromosome is a single piece of DNA that contains many genes, regulatory sequence and other genetic sequence....
s and are the mechanical basis of heredity. These discoveries formed the basis of the modern science of genetics
Genetics

Genetics , a discipline of biology, is the science of heredity and Genetic variation in living organisms. The fact that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding....
. When he was awarded the Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine

The Nobel Prize in Physiology or Medicine is awarded once a year by the Swedish Karolinska Institutet. It is one of the five Nobel Prizes established by the will of Alfred Nobel in 1895, awarded for outstanding contributions in Nobel Prize in Physics, Nobel Prize in Chemistry, Nobel Prize in Literature, Nobel Peace Prize, and Physiology or Medic...
 in 1933 he was the first person awarded the Prize in genetics, for his discoveries concerning the role played by the chromosome in heredity.

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
Model organism

A model organism is a species that is extensively studied to understand particular biology phenomena, with the expectation that discoveries made in the organism model will provide insight into the workings of other organisms....
 in contemporary genetics. The Division of Biology he established at the California Institute of Technology
California Institute of Technology

The California Institute of Technology is a private university research university located in Pasadena, California, United States. Caltech maintains a strong emphasis on the natural sciences and engineering....
 produced seven Nobel Prize winners.

Early life

len Key Howard Morgan. Part of a long line of Southern
Southern United States

The Southern United States—commonly referred to as the American South, Dixie, or simply the South—constitutes a large distinctive region in the southeastern and south-central United States....
 aristocracy on his father's side, Morgan was a nephew of Confederate
Confederate States Army

The Confederate States Army was a military organization whose primary mission was to provide the necessary forces and capabilities to support the National Security and defense of the Confederate States of America during its brief existence from 1861 to 1865....
 General John Hunt Morgan
John Hunt Morgan

John Hunt Morgan was a Confederate States Army General officer and cavalry officer in the American Civil War.Morgan is best known for Morgan's Raid in 1863, when he led 2,460 troops racing past Union Army lines into Kentucky, Indiana, and Ohio in July 1863....
 and his great-grandfather John Wesley Hunt
John Wesley Hunt

John Wesley Hunt was a prominent businessman and early civic leader in Lexington, Kentucky. He was one of the first millionaires west of the Allegheny Mountains....
 had been the first millionaire west of the Allegheny Mountains
Allegheny Mountains

The Allegheny Mountain Range — informally, the Alleghenies — is part of the vast Appalachian Mountains of the eastern United States and Canada....
. Through his mother, he was the great-grandson of Francis Scott Key
Francis Scott Key

Francis Scott Key was an United States lawyer, author, and amateur poet, from Georgetown, Washington, D.C., who wrote the words to the United States' national anthem, "The Star-Spangled Banner."...
, the author of the "Star Spangled Banner", and John Eager Howard
John Eager Howard

John Eager Howard was an United States soldier and politician from Maryland. He was born in and died in Baltimore County, Maryland. Howard County, Maryland, is named for him....
, a one-time governor and senator from Maryland
Maryland

Maryland is a U.S. state located in the Mid Atlantic States of the United States, bordering Virginia, West Virginia and the Washington, D.C. to the south and west, Pennsylvania to the north, and Delaware to the east....
. 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
University of Kentucky

The University of Kentucky, also known as UK, is a state university , co-educational, university, and is also the state's land-grant university, located in Lexington, Kentucky, Kentucky....
). There, he focused on science; he particularly enjoyed natural history, and worked with the U.S. Geological Survey
United States Geological Survey

The United States Geological Survey is a scientific agency of the United States government. The scientists of the USGS study the landscape of the United States, its natural resources, and the natural hazards that threaten it....
 in his summers. He graduated as valedictorian
Valedictorian

Valedictorian is an academic title typically conferred in North America upon the highest ranked student among those being graduated from an educational institution....
 in 1886 and was the only student to graduate with a bachelor in science. Following a summer at the Marine Biology School in Annisquam, Massachusetts
Annisquam, Massachusetts

Annisquam is a small neighborhood on the northern coast of Massachusetts, and a neighborhood of the city of Gloucester, Massachusetts.A few miles from downtown Gloucester, Annisquam is primarily a residential neighborhood, and Annisquam's only businesses include a restaurant and marina, a small hotel, a Real Estate Company, a library and...
, Morgan began graduate studies in zoology
Zoology

Zoology is the branch of biology concerned with the study of animals. The most common pronunciation of "zoology" is ; however, an alternative pronunciation is ....
 at the recently founded Johns Hopkins University
Johns Hopkins University

The Johns Hopkins University, commonly referred to as Hopkins or JHU, is a private university research university located in Baltimore, Maryland, Maryland, United States....
, the first research-oriented American university. After two years of experimental work with morphologist
Morphology (biology)

The term morphology in biology refers to form, structure and configuration of an organism. This includes aspects of the outward appearance as well as the form and structure of the internal parts like bones and organs....
 William Keith Brooks
William Keith Brooks

William Keith Brooks, LL.D., Ph.D. was an United States zoologist, born in Cleveland, Ohio, March 25, 1848. He was educated at Williams College and at Harvard University ....
 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 spider
Sea spider

Sea spiders, also called Pantopoda or pycnogonids , are marine arthropods of class Pycnogonida. They are cosmopolitan distribution, found especially in the Mediterranean Sea and Caribbean Seas and the Arctic Ocean and Antarctic Oceans....
s—collected during the summers of 1889 and 1890 at the Marine Biological Laboratory
Marine Biological Laboratory

The Marine Biological Laboratory is an international center for research and education in biology and ecology. Founded in 1888, the MBL is the oldest independent marine laboratory in the Americas, taking advantage of a coastal setting in the Cape Cod village of Woods Hole, Massachusetts....
 in Woods Hole, Massachusetts
Woods Hole, Massachusetts

Woods Hole is a census-designated place in the town of Falmouth , Massachusetts in Barnstable County, Massachusetts, Massachusetts, United States....
—to determine their phylogenetic
Phylogenetics

In biology, phylogenetics is the study of evolutionary relatedness among various groups of organisms , which is discovered through molecular sequencing data and morphological data matrices....
 relationship with other arthropod
Arthropod

Arthropods are animals belonging to the Scientific classification Arthropoda , and include the insects, arachnids, crustaceans, and others....
s. He concluded that with respect to embryology they were more closely related to spiders
Chelicerata

The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites....
 than crustaceans. Based on the publication of this work Morgan was awarded his Ph.D.
Doctor of Philosophy

Doctor of Philosophy, abbreviated Ph.D. or PhD for the Latin , meaning "teacher of philosophy", is an postgraduate academic degree awarded by University....
 from Johns Hopkins in 1890, and was also awarded the Bruce Fellowship in Research. With the fellowship he was able to travel to Jamaica
Jamaica

Jamaica is an island nation of the Greater Antilles, in length and as much as in width situated in the Caribbean Sea. It is about south of Cuba, and west of the island of Hispaniola, on which Haiti and the Dominican Republic are situated....
, the Bahamas and to Europe
Europe

Europe is, conventionally, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally divided from Asia to its east by the water divide of the Ural Mountains, the Ural , the Caspian Sea, and by the Caucasus Mountains to the southeast....
 to conduct further research.

Bryn Mawr

In 1891, Morgan was appointed associate professor (and head of the biology department) at Johns Hopkins' sister school Bryn Mawr College
Bryn Mawr College

'Bryn Mawr College' is a highly selective Women's colleges in the United States Liberal arts colleges in the United States located in Bryn Mawr, Pennsylvania, a community in Lower Merion Township, Pennsylvania, Pennsylvania, ten miles west of Philadelphia, Pennsylvania....
, replacing his colleague E. B. Wilson
Edmund Beecher Wilson

Edmund Beecher Wilson was a pioneering United States zoologist and geneticist.Wilson was born in Geneva, Illinois, Illinois, and graduated from Yale University in 1878....
. Morgan taught all morphology-related courses, while the other member of the department, Jacques Loeb
Jacques Loeb

Jacques Loeb was a Germany-born United States of America physiologist and biologist....
, 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.

Stazionezoologica
In 1894 Morgan was granted a year's absence to conduct research in the laboratories of Stazione Zoologica
Stazione Zoologica

The Stazione Zoologica Anton Dohrn is a research institute in Naples, Italy, devoted to basic research in biology. Research is largely interdisciplinary involving the fields of evolution, biochemistry, molecular biology, neurobiology, cell biology, biological oceanography, marine botany, molecular plant biology, benthic ecology, and ecophysi...
 in Naples
Naples

Naples is a city in southern Italy, the capital of the region of Campania and of the province of Naples. The city is known for its rich history, art, culture and gastronomy, playing an important role throughout much of its existence; it is over 2,800 years old....
, 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
Ctenophore

The Ctenophora , commonly known as comb jellies, is a phylum of animals that live in all types of marine waters world-wide. Their most distinctive feature is the "combs", groups of cilia that they use for swimming, and they are the largest animals that swim by means of cilia ? adults of various species range from a few millimeters to...
 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
Naturphilosophie

Naturphilosophie was a current in the philosophy tradition of German idealism in the 19th century, particularly associated with Friedrich Wilhelm Joseph von Schelling and Georg Wilhelm Friedrich Hegel....
 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
Wilhelm Roux

Wilhelm Roux was a Germany zoology and pioneer of experimental embryology.Roux was born and educated in Jena, Germany where he attended university and studied under Ernst Haeckel....
'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 protoplasam 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
Sea urchin

Sea urchins are small, spiny, globular creatures that compose most of class Echinoidea. They are found in oceans all over the world. Their shell, or "test", is round and spiny, typically from 3 to 10 cm across....
 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
Sea urchin

Sea urchins are small, spiny, globular creatures that compose most of class Echinoidea. They are found in oceans all over the world. Their shell, or "test", is round and spiny, typically from 3 to 10 cm across....
 eggs could be induced to divide without fertilization by adding magnesium chloride
Magnesium chloride

Magnesium chloride is the name for the chemical compounds with the chemical formulas MgCl2 and its various water of hydrations MgCl2x....
, 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
Regeneration (biology)

In biology, an organism is said to regenerate a lost or damaged part if the part regrows so that the original function is restored.Regenerative capacity is inversely related to complexity: in general, the more complex an animal is the less regeneration it is capable of....
 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 was 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
Lilian Vaughn Morgan

Lilian Vaughn Morgan was an American genetics and the wife of Thomas Hunt Morgan. Ms. Morgan published 16 single author papers in her lifetime....
 (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
Columbia University

Columbia University in the City of New York , is a private university in the United States and a member of the Ivy League. Columbia's main campus lies in the Morningside Heights, Manhattan neighborhood in the borough of Manhattan, in New York City....
, which at last freed him to focus fully on experimental work.
One of their four children (one boy and three girls) was Isabel Morgan
Isabel Morgan

Isabel Merrick Morgan was an American virologist at Johns Hopkins University,who – in a research team with David Bodian and Howard Howe – prepared an experimental vaccine that protected monkeys against poliomyelitis....
 (1911–1996) (marr. Mountain), who became a virologist at Johns Hopkins, specializing in polio
Poliomyelitis

Poliomyelitis, often called polio or infantile paralysis, is an acute virus infectious disease spread from person to person, primarily via the fecal-oral route....
 research.

Columbia University

By 1904, when Morgan took a professorship in experimental zoology at Columbia University
Columbia University

Columbia University in the City of New York , is a private university in the United States and a member of the Ivy League. Columbia's main campus lies in the Morningside Heights, Manhattan neighborhood in the borough of Manhattan, in New York City....
, he was becoming increasingly focused on the mechanisms of heredity and evolution. The previous year, he had published Evolution and Adaption; like many biologists at that time, he saw clear evidence for biological evolution (as in the common descent
Common descent

A group of organisms is said to have common descent if they have a common ancestor. In modern biology, it is generally accepted that all living organisms on Earth are descended from a common ancestor or ancestral gene pool....
 of similar species) but rejected Darwin's proposed mechanism of natural selection
Natural selection

Natural selection is the process by which favorable heritable trait become more common in successive generations of a population of Reproduction organisms, and unfavorable heritable traits become less common, due to differential reproduction of genotypes....
 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 Larmarckian
Lamarckism

Lamarckism is the once widely accepted idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring ....
 mechanism of inheritance of acquired characters
Inheritance of acquired characters

The inheritance of acquired traits is a hypothesis about a mechanism of heredity by which changes in physiology acquired over the life of an organism may purportedly be transmitted to offspring....
, which featured prominently in Darwin's theory, was increasingly rejected by biologists. According to Morgan 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
Carl Correns

Carl Erich Correns was a Germany botanist and geneticist, who is notable primarily for his independent discovery of the principles of heredity, and for his rediscovery of Gregor Mendel's earlier paper on that subject, which he achieved simultaneously but independently of the botanists Erich Tschermak von Seysenegg and Hugo de Vries....
, Erich von Tschermak
Erich von Tschermak

Erich von Tschermak-Seysenegg was an Austrian agronomist.von Tschermak is one of three men - see also Hugo de Vries and Carl Correns - who independently rediscovered Gregor Mendel's work on genetics....
 and Hugo De Vries
Hugo de Vries

Hugo Marie de Vries was a Netherlands botanist and one of the first geneticists. He is known chiefly for suggesting the concept of genes, rediscovering Gregor Mendel's laws of heredity in the 1890s, and for developing a mutation theory of evolution....
 had rediscovered the work of Gregor Mendel
Gregor Mendel

Gregor Johann Mendel was an Augustinians priest and scientist, and is often called the father of genetics for his study of the biological inheritance of certain Trait s in pea plants....
, and with it the foundation of genetics
Genetics

Genetics , a discipline of biology, is the science of heredity and Genetic variation in living organisms. The fact that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding....
. 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
Hugo de Vries

Hugo Marie de Vries was a Netherlands botanist and one of the first geneticists. He is known chiefly for suggesting the concept of genes, rediscovering Gregor Mendel's laws of heredity in the 1890s, and for developing a mutation theory of evolution....
' 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.

Sexlinked Inheritance White
Following C. W. Woodworth
Charles W. Woodworth

Charles W. Woodworth was an American Entomology. He founded the Entomology Department at the University of California, Berkeley, and made many valuable contributions to entomology during his career....
 and William E. Castle
William E. Castle

William Ernest Castle was an early United States genetics....
, around 1908 Morgan started working on the fruit fly Drosophila melanogaster
Drosophila melanogaster

Drosophila melanogaster is a two-winged insect that belongs to the Diptera, the Order of the Fly. The species is commonly known as the Drosophilidae or vinegar fly, and is one of the most commonly used model organisms in biology, including studies in genetics, physiology and Life history theory....
, and encouraging students to do so as well. With Fernandus Payne
Fernandus Payne

Fernandus Payne was an United States zoologist, geneticist and educator.Panye was born in Shelbyville, Indiana. He received a B.Sc. from Valparaiso University in 1901 and a B.A....
, 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
Mutant

A mutant is an individual, organism, or new genetic character arising or resulting from an instance of mutation, which is a base-pair sequence change within the DNA of a gene or chromosome of an organism resulting in the creation of a new character or Trait not found in the wild type....
 male among the red-eyed wild type
Wild type

Wild type, sometimes written wildtype or wild-type, is the typical form of an organism, strain, gene, or characteristic as it occurs in nature....
s. 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
White (mutation)

white was the first sex-linked mutation ever discovered in Drosophila melanogaster. In 1910 Thomas Hunt Morgan, collected a single male white-eyed mutant from a population of Drosophila melanogaster fruit flies, which usually have bright red eyes....
. Morgan also discovered a pink-eyed mutant that showed a different pattern of inheritance. In a paper published in Science
Science (journal)

Science is the academic journal of the American Association for the Advancement of Science and is considered one of the world's most prestigious scientific journals....
 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 Crossover 1
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
Genetic linkage

Genetic linkage occurs when particular genetic Locus or alleles for genes are inherited jointly. Genetic loci on the same chromosome are physically connected and tend to stay together during meiosis, and are thus genetically linked....
 and to hypothesize the phenomenon of crossing over
Crossing Over

Crossing Over may refer to:* Chromosomal crossover, a cellular process* Crossing Over , a 1998 album by Hesperus * Crossing Over , a book by John Edward...
. 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
J. B. S. Haldane

John Burdon Sanderson Haldane Royal Society#Fellowship , known as Jack , was a UK-born geneticist and evolutionary biologist. He was one of the founders of population genetics....
 suggested that the unit of measurement for linkage be called the morgan
Centimorgan

In genetics, a centimorgan or map unit is a unit of recombinant frequency for measuring genetic linkage. It is often used to imply distance along a chromosome....
. Morgan's student Alfred Sturtevant
Alfred Sturtevant

Alfred Henry Sturtevant was an United States geneticist. Sturtevant constructed the first genetic map of a chromosome in 1913. Throughout his career he worked on the organism Drosophila melanogaster with Thomas Hunt Morgan....
 developed the first genetic map in 1913.

In 1915 Morgan, Sturtevant, Calvin Bridges
Calvin Bridges

Calvin Blackman Bridges was an American scientist, known for his contributions to the field of genetics. Along with Alfred Sturtevant and H.J. Muller, Bridges was part of the famous fly room of Thomas Hunt Morgan at Columbia University....
 and H. J. Muller wrote the seminal book The Mechanism of Mendelian Heredity. Geneticist Curt Stern
Curt Stern

Curt Stern was a Germany-United States geneticist. He made several important genetic discoveries, demonstrating chromosomal crossover in Drosophila weeks after Barbara McClintock and Harriet Creighton had done so [with maize] in 1931....
 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" pioneered by Morgan and his students. Garland Allen characterized the post-1915 period as one of normal science
Normal science

Normal science is a concept originated by Thomas Samuel Kuhn and elaborated in The Structure of Scientific Revolutions. The term refers to the relatively routine work of scientists experimenting within a paradigm, slowly accumulating detail in accord with established broad theory, not actually challenging or attempting to test the underly...
, 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
Gene

A gene is the basic unit of heredity in a living organism. All living things depend on genes. Genes hold the information to build and maintain their cell and pass genetic trait to offspring....
 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
Richard Goldschmidt

Richard Benedict Goldschmidt was a Germany-born United States geneticist. He is considered the first to integrate genetics, development, and evolution....
 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 organism
Model organism

A model organism is a species that is extensively studied to understand particular biology phenomena, with the expectation that discoveries made in the organism model will provide insight into the workings of other organisms....
s. 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
Eugenics

Eugenics is a scientific field involving the controlled breeding of humans in order to achieve desirable traits in future generations. Eugenics was at its height in first half of the 20th century and was largely abandoned with the end of World War II....
 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
United States National Academy of Sciences

The National Academy of Sciences is a corporation in the United States whose members serve pro bono as "advisers to the nation on science, engineering, and medicine."...
 and made a foreign member of the Royal Society
Royal Society

The Royal Society of London for the Improvement of Natural Knowledge, known simply as the Royal Society, or even the Royal, is a learned society for science that was founded in 1660 and is considered by most to be the oldest such society still in existence....
. In 1924 Morgan received the Darwin Medal
Darwin Medal

The Darwin Medal is awarded by the Royal Society every alternate year for "work of acknowledged distinction in the broad area of biology in which Charles Darwin worked"....
. 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
George Ellery Hale

George Ellery Hale was an American Sun astronomer, born in Chicago, Illinois. He was educated at Massachusetts Institute of Technology, at the Observatory of Harvard College, , and at Humboldt University of Berlin ....
 to establish a school of biology in California.

Caltech

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

The California Institute of Technology is a private university research university located in Pasadena, California, United States. Caltech maintains a strong emphasis on the natural sciences and engineering....
 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 on 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
Albert Tyler (biologist)

Albert Tyler was an United States biology whose research was focused on reproductive biology and development in marine organisms.Tyler was born in Brooklyn, New York....
 from Columbia and took on Theodosius Dobzhansky
Theodosius Dobzhansky

Theodosius Grygorovych Dobzhansky, also known as T. G. Dobzhansky, and sometimes Anglicized to Theodore Dobzhansky was a noted genetics and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis....
 as an international research fellow. More scientists came to work in the Division including George Beadle, Boris Ephrussi
Boris Ephrussi

Boris Ephrussi was a France genetics of Russian origin. He was one of the many famous Jewish Biology. He had published two papers in November 1966 which represented a key step in a decade of research in his laboratory....
, Edward L. Tatum, Linus Pauling
Linus Pauling

Linus Carl Pauling was an United States scientist, peace activist, author and list of educators. He was one of the most influential chemists in history and ranks among the most important scientists in any field of the 20th century....
, Frits Went
Frits Went

Friedrich August Ferdinand Christian Went was a Netherlands botanist, professor of botany and director of the Botanical Garden at the University of Utrecht....
, and Sidney W. Fox
Sidney W. Fox

Sidney Walter Fox was a Los Angeles-born biochemistry responsible for unique discoveries in the autosynthesis of protocells....
.

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
American Association for the Advancement of Science

The American Association for the Advancement of Science is an international non-profit organization with the stated goals of promoting cooperation between scientists, defending scientific freedom, encouraging scientific responsibility, and supporting science education and science outreach for the betterment of all humanity....
; and in 1932 he chaired the Sixth International Congress of Genetics in Ithaca, New York
Ithaca, New York

The City of Ithaca sits on the southern shore of Cayuga Lake, in Central New York New York State, USA. It is best known for being home to Cornell University ? an Ivy League school with almost 20,000 students ....
. In 1933 Morgan was awarded the Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine

The Nobel Prize in Physiology or Medicine is awarded once a year by the Swedish Karolinska Institutet. It is one of the five Nobel Prizes established by the will of Alfred Nobel in 1895, awarded for outstanding contributions in Nobel Prize in Physics, Nobel Prize in Chemistry, Nobel Prize in Literature, Nobel Peace Prize, and Physiology or Medic...
; 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 chromosome
Polytene chromosome

To increase Cell volume, some specialized cells undergo repeated rounds of DNA replication without cell division , forming a giant polytene chromosome....
s 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
Copley Medal

The Copley Medal is an award given by the Royal Society of London for "outstanding achievements in research in any branch of science, and alternates between the physical sciences and the biological sciences"....
 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.

Legacy

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
George Wells Beadle

George Wells Beadle was an United States scientist in the field of genetics, and Nobel Prize in Physiology or Medicine Nobel laureate who with Edward Lawrie Tatum discovered the role of genes in regulating biochemical events within cells....
, Edward B. Lewis
Edward B. Lewis

Edward B. Lewis was an United States geneticist, a corecipient of the 1995 Nobel Prize in Physiology or Medicine.Lewis was born in Wilkes-Barre, Pennsylvania and graduated from E.L....
 and Hermann Joseph Muller
Hermann Joseph Muller

Hermann Joseph ?H.J.? Muller was an United States geneticist, educator, and Nobel laureate best known for his work on the physiological and genetic effects of radiation as well as his outspoken political beliefs....
. 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
Genetics Society of America

The Genetics Society of America is a non-exclusive association of genetics researchers and educators, and the publisher of the peer-reviewed journal Genetics ....
 annually awards the Thomas Hunt Morgan Medal
Thomas Hunt Morgan Medal

The Thomas Hunt Morgan Medal is a medal awarded for lifetime contributions to the field of genetics. The medal is awarded by the Genetics Society of America....
 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.

See also

  • List of books by Thomas Hunt Morgan
    List of books by Thomas Hunt Morgan

    This is a list of books and monographs by the American geneticist Thomas Hunt Morgan. Morgan produced 22 books on embryology, genetics and evolution....
  • History of genetics
    History of genetics

    The history of genetics is generally held to have started with the work of an Augustinian monk, Gregor Mendel. Experiments on Plant Hybridization on pea plants, published in 1866, described what came to be known as Mendelian Inheritance....
  • History of model organisms
    History of model organisms

    The history of model organisms began with the idea that certain organisms can be studied and used to gain knowledge of other organisms or as a control for other organisms of the same species....
  • Centimorgan
    Centimorgan

    In genetics, a centimorgan or map unit is a unit of recombinant frequency for measuring genetic linkage. It is often used to imply distance along a chromosome....
     - unit of genetic crossover


Further Reading



External links