Hermann Joseph Muller (December 21, 1890 – April
5, 1967) was an American geneticist, educator,
and Nobel laureate best known for his work
on the physiological and genetic effects of
radiation (mutagenesis) as well as his outspoken
political beliefs. Muller frequently warned
of long-term dangers of radioactive fallout
from nuclear war and nuclear testing, which
resulted in greater public scrutiny of these
practices.
== Early life ==
Muller was born in New York City, the son
of Frances (Lyons) and Hermann Joseph Muller,
Sr., an artisan who worked with metals. Muller
was a third-generation American whose
father's ancestors were originally Catholic
and came to the United States from Koblenz.
His mother's family was of mixed Jewish (descended
from Spanish and Portuguese Jews) and Anglican
background, and had come from Britain. Among
his first cousins are Herbert J. Muller and
Alfred Kroeber whose daughter is Ursula Le
Guin. As an adolescent, he attended a Unitarian
church and considered himself a pantheist;
in high school he became an atheist. He excelled
in the public schools. At 16 he entered Columbia
College. From his first semester he was interested
in biology; he became an early convert of
the Mendelian-chromosome theory of heredity
— and the concept of genetic mutations and
natural selection as the basis for evolution.
He formed a Biology Club and also became a
proponent of eugenics; the connections between
biology and society would be his perennial
concern. Muller earned a B.A. degree in 1910.Muller
remained at Columbia (the pre-eminent American
zoology program at the time, thanks to E.
B. Wilson and his students) for graduate school.
He became interested in the Drosophila genetics
work of Thomas Hunt Morgan's fly lab after
undergraduate bottle washers Alfred Sturtevant
and Calvin Bridges joined his Biology Club.
In 1911-1912, he studied metabolism at Cornell
University, but remained involved with Columbia.
He followed the drosophilists as the first
genetic maps emerged from Morgan's experiments,
and joined Morgan's group in 1912 (after two
years of informal participation).In the fly
group, Muller's contributions were primarily
theoretical: explanations for experimental
results and ideas and predictions for new
experiments. In the emerging collaborative
culture of the drosophilists, however, credit
was assigned based on results rather than
ideas; Muller felt cheated when he was left
out of major publications.
== Career ==
In 1914, Julian Huxley offered Muller a position
at the recently founded William Marsh Rice
Institute, now Rice University; he hurried
to complete his Ph.D. degree and moved to
Houston for the beginning of the 1915-1916
academic year (his degree was issued in 1916).
At Rice, Muller taught biology and continued
Drosophila lab work. In 1918, he proposed
an explanation for the dramatic discontinuous
alterations in Oenothera larmarckiana that
were the basis of Hugo de Vries's
theory of mutationism: "balanced lethals"
allowed the accumulation of recessive mutations,
and rare crossing over events resulted in
the sudden expression of these hidden traits.
In other words, de Vries's experiments were
explainable by the Mendelian-chromosome theory.
Muller's work was increasingly focused on
mutation rate and lethal mutations. In 1918,
Morgan—short-handed because many of his
students and assistants were drafted for the
U.S. entry into World War I—convinced Muller
to return to Columbia to teach and to expand
his experimental program.At Columbia, Muller
and his collaborator and longtime friend Edgar
Altenburg continued the investigation of lethal
mutations. The primary method for detecting
such mutations was to measure the sex ratios
of the offspring of female flies. They predicted
the ratio would vary from 1:1 due to recessive
mutations on the X chromosome, which would
be expressed only in males (who lacked the
functional allele on a second X chromosome).
Muller found a strong temperature dependence
in mutation rate, leading him to believe that
spontaneous mutation was the dominant mode
(and to initially discount the role of external
factors such as ionizing radiation or chemical
agents). In 1920, Muller and Altenburg coauthored
a seminal paper in Genetics on "modifier genes"
that determine the size of mutant Drosophila
wings. In 1919, Muller made the important
discovery of a mutant (later found to be a
chromosomal inversion) that appeared to suppress
crossing-over, which opened up new avenues
in mutation rate studies. However, his appointment
at Columbia was not continued; he accepted
an offer from the University of Texas and
left Columbia after the summer of 1920.Muller
taught at The University of Texas from 1920
until 1932. Soon after returning to Texas,
he married mathematics professor Jessie Marie
Jacobs, whom he had courted previously. In
his early years at Texas, Muller's Drosophila
work was slow going; the data from his mutation
rate studies were difficult to interpret.
In 1923, he began using radium and X-rays,
but the relationship between radiation and
mutation was difficult to measure because
such radiation also sterilized the flies.
In this period, he also became involved with
eugenics and human genetics. He carried out
a study of twins separated at birth that seemed
to indicate a strong hereditary component
to I.Q. Muller was critical of the new directions
of the eugenics movement (such as anti-immigration),
but was hopeful about the prospects for positive
eugenics. In 1932, at the Third International
Eugenics Congress, Muller gave a speech and
stated "eugenics might yet perfect the human
race but only in a society consciously organized
for the common good.
=== Discovery of X-ray mutagenesis ===
1926 marked the beginning of a series of major
breakthroughs. Beginning in November, Muller
carried out two experiments with varied doses
of X-rays, the second of which used the crossing
over suppressor stock ("ClB") he had found
in 1919. A clear, quantitative connection
between radiation and lethal mutations quickly
emerged. Muller's discovery created a media
sensation after he delivered a paper entitled
"The Problem of Genetic Modification" at the
Fifth International Congress of Genetics in
Berlin; it would make him one of the better
known public intellectuals of the early 20th
century. By 1928, others had replicated his
dramatic results, expanding them to other
model organisms such as wasps and maize. In
the following years, he began publicizing
the likely dangers of radiation exposure in
humans (such as physicians who frequently
operate X-ray equipment).His lab grew quickly,
but it shrank again following the onset of
the Great Depression. Especially after the
stock market crash, Muller was increasingly
pessimistic about the prospects of capitalism.
Some of his visiting lab members were from
the USSR, and he helped edit and distribute
an illegal leftist student newspaper, The
Spark. It was a difficult period for Muller
both scientifically and personally: his marriage
was falling apart, and he was increasingly
dissatisfied with his life in Texas. Meanwhile,
the waning of the eugenics movement, ironically
hastened by his own work pointing to the previously
ignored connections between environment and
genetics, meant that his ideas on the future
of human evolution had reduced impact in the
public sphere.
=== Work in Europe ===
In September 1932, Muller moved to Berlin
to work with the Russian expatriate geneticist
Nikolay Timofeeff-Ressovsky, a trip intended
as a limited sabbatical stretched into an
eight-year, five-country journey. In Berlin,
he met two physicists who would later be significant
to the biology community: Niels Bohr and Max
Delbrück. The Nazi movement was precipitating
the rapid emigration of scientific talent
from Germany, and Muller was particularly
opposed to the politics of National Socialism.
But the FBI was investigating Muller because
of his involvement with The Spark, so he chose
instead to go to the Soviet Union (an environment
better suited to his political beliefs). In
1933, Muller and his wife reconciled, and
she and their son David E. Muller moved with
Hermann to Leningrad. There, at the Institute
of Genetics, he imported the basic equipment
for a Drosophila lab—including the flies—and
set up shop. The Institute was moved to Moscow
in 1934, and Muller and his wife were divorced
in 1935.In the USSR, Muller supervised a large
and productive lab, and organized work on
medical genetics. Most of his work involved
further explorations of genetics and radiation.
There he completed his eugenics book, Out
of the Night, the main ideas of which dated
to 1910. By 1936, however, Joseph Stalin's
repressive policies and the rise of Lysenkoism
was making the USSR an increasingly problematic
place to live and work. Muller and much of
the Russian genetics community did what they
could to oppose Trofim Lysenko and his Larmarckian
evolutionary theory, but Muller was soon forced
to leave the Soviet Union after Stalin read
a translation of his eugenics book and was
"displeased by it, and...ordered an attack
prepared against it."Muller—with about 250
strains of Drosophila—moved to Edinburgh
in September 1937, after a brief stay in Madrid
and Paris. In 1938, with war on the horizon,
he began looking for a permanent position
back in the United States. He also began courting
Dorothea "Thea" Kantorowicz, a German refugee;
they were married in May 1939. The Seventh
International Congress on Genetics was held
in Edinburgh later that year; Muller wrote
a "Geneticists' Manifesto" in response to
the question: "How could the world's population
be improved most effectively genetically?"
He also engaged in a debate with the perennial
genetics gadfly Richard Goldschmidt over the
existence of the gene, for which there remained
little direct physical evidence.
=== Later career ===
When Muller returned to the United States
in 1940, he took an untenured research position
at Amherst College, in the department of Otto
C. Glaser. After the U.S. entry into World
War II, his position was extended indefinitely
and expanded to include teaching. His Drosophila
work in this period focused on measuring the
rate of spontaneous (as opposed to radiation-induced)
mutations. Muller's publication rate decreased
greatly in this period, from a combination
of lack of lab workers and experimentally
challenging projects. However, he also worked
as an adviser in the Manhattan Project (though
he did not know that was what it was), as
well as a study of the mutational effects
of radar. Muller's appointment was ended after
the 1944–1945 academic year, and despite
difficulties stemming from his socialist political
activities, he found a position as professor
of zoology at Indiana University. Here, he
lived in a Dutch Colonial Revival house in
Bloomington's Vinegar Hill neighborhood.In
1946 Muller was awarded the Nobel Prize in
Physiology or Medicine, "for the discovery
that mutations can be induced by x-rays".
Genetics, and especially the physical and
physiological nature of the gene, was becoming
a central topic in biology, and X-ray mutagenesis
was a key to many recent advances, among them
George Beadle and Edward Tatum's work on Neurospora
that established in 1941 the one gene-one
enzyme hypothesis. In Muller's Nobel Prize
lecture he argued that there was no threshold
dose of radiation that did not produce mutagenesis,
which led to the adoption of the linear no-threshold
model of radiation on cancer risks.The Nobel
Prize, in the wake of the atomic bombings
of Hiroshima and Nagasaki, focused public
attention on a subject Muller had been publicizing
for two decades: the dangers of radiation.
In 1952, nuclear fallout became a public issue;
since Operation Crossroads, more and more
evidence had been leaking out about radiation
sickness and death caused by nuclear testing.
Muller—and many other scientists—pursued
an array of political activities to defuse
the threat of nuclear war. With the Castle
Bravo fallout controversy in 1954, the issue
became even more urgent. In 1955 Muller was
one of eleven prominent intellectuals to sign
the Russell-Einstein Manifesto, the upshot
of which was the first 1957 Pugwash Conference
on Science and World Affairs, which addressed
the control of nuclear weapons. He was a signatory
(with many other scientists) of the 1958 petition
to the United Nations, calling for an end
to nuclear weapons testing, which was initiated
by the Nobel Prize-winning chemist Linus Pauling.It
has however been argued that Muller's opinions
on the effect of radiation on mutagenesis
were not supported by studies on the survivors
of the atomic bombings, or in research on
mice. Some scientists have criticized the
approach of Muller; Geneticist James F. Crow
called Muller's view "alarmist" and wrote
that it created in the public "an irrational
fear of low-level radiation relative to other
risks". It is also argued that he was familiar
with another study that directly contradicted
the linear no-threshold model he supported,
leading to a charge in 2011 that Muller misled
the public as to the then-current data available
and that would have an effect on the formulation
of policy that favored his model.Muller was
awarded the Linnean Society of London's Darwin-Wallace
Medal in 1958 and the Kimber Genetics Award
of the U.S. National Academy of Sciences in
1955. He served as president of the American
Humanist Association from 1956 to 1958. The
American Mathematical Society selected him
as its Gibbs Lecturer for 1958. He retired
in 1964.H. J. Muller and science fiction writer
Ursula Le Guin were second cousins; his father
(Hermann J. Muller Sr.) and her father's mother
(Johanna Muller Kroeber) were siblings, the
children of Nicholas Müller, who immigrated
to the United States in 1848, and at that
time dropped the umlaut from his name. Another
cousin was Herbert J. Muller, whose grandfather
Otto was another son of Nicholas and a sibling
of Hermann Sr. and Johanna.
== Personal life ==
Muller is survived by his daughter, Helen
J. Muller, now a professor emerita at the
University of New Mexico, who has a daughter,
Mala Htun. His son, David E. Muller, a professor
emeritus of mathematics and computer science
at the University of Illinois and at New Mexico
State University, died in 2008 in Las Cruces,
New Mexico. David's mother was Jessie Jacobs
Muller Offermann, Hermann's first wife. Helen's
mother was Dorothea Kantorowicz Muller, Hermann's
second wife.
== Former graduate students ==
Seymour Abrahamson
Raissa L. Berg
Elof Axel Carlson
Sara Helen Frye
H. Bentley Glass
C. P. Oliver
Irwin I. Oster
Abraham P. Schalet
Wilson Stone
William Edgar Trout III
Dale Eugene WagonerFormer post-doctoral fellowsGeorge
D. SnellWorked in lab as undergraduatesMargaret
Russell Edmondson
Carl SaganPeople who worked in his lab in
Indiana [1]
== Bibliography ==
Herman Joseph Muller, Modern Concept of Nature
(SUNY Press, 1973). ISBN 0-87395-096-8.
Herman Joseph Muller, Man's Future Birthright
(SUNY Press, 1973). ISBN 0-87395-097-6.
H. J. Muller, Out of the Night: A Biologist's
View of the Future (Vanguard Press, 1935).
H. J. Muller, Studies in Genetics: The Selected
Papers of H. J. Muller (Indiana University
Press, 1962).
== See also ==
Mutagenesis
Bateson–Dobzhansky–Muller model
Repository for Germinal Choice
Muller's ratchet
Muller's morphs
History of biology
History of genetics
History of model organisms
== References ==
== External links ==
Nobel Biography
Hermann Joseph Muller — Biographical Memoirs
of the National Academy of Sciences
The Muller manuscripts, 1910–1967 in archives
of the Indiana University
On the origins of the linear no-threshold
(LNT) dogma by means of untruths, artful dodges
and blind faith, Edward J. Calabrese, Environmental
Research 142 (2015) 432–442.
Hermann J. Muller Collection Cold Spring Harbor
Laboratory Archives
