Hermann Joseph Muller was a professor, geneticist, and Nobel laureate who is best known as the founder of the field of radiation genetics, in addition to being a co-founder of the American school of classical genetics. Much of the research that led Muller to the Nobel prize was done at UT Austin in the Biological Sciences building (BIO) during the 1920s. But for all the accolades Muller received for his work in genetics, he lived an intense personal and professional life and found himself in many political hotbeds of the world during his lifetime.

Muller identified as both an atheist and a humanist, and while an advocate of selective breeding for humanity, he was a vocal opponent of the American eugenics movement in the early 20th century, accusing it of racism and elitism. In addition, Muller was committed to Darwinism and supported natural selection as the basic mechanism of evolution, a view under attack in academia in the early part of the 20th century. His communist-leaning political beliefs also lead him into trouble with UT and the FBI, and ultimately drew him to Europe during World War II, where he saw the rise of Nazism and collided with Stalinism. In addition, his first marriage suffered immensely from the strain of his intense work hours and his outspoken beliefs. Nevertheless, Muller retained the respect of geneticists the world over.

MULLER'S EARLY LIFE

Muller was born in New York City on December 21, 1890, to Hermann Joseph and Frances Lyons, both first-generation Americans. Hermann Joseph continued his own father's fine art metal business but sadly died of a cerebral stroke when young Hermann was only ten.

As a child, Muller displayed an early interest in science. It is believed he and his friends started what might have been the first high school science club in the US.

Muller would later earn his bachelors in zoology in 1910 at Columbia University but had the additional responsibility of supporting his mother and sister since his father had passed on. To help his family economically, Muller worked odd jobs while in school, serving as a runner on Wall Street and an English teacher to immigrants. While working towards his Masters in nerve physiology at Cornell, he taught physiology for a year at Cornell University Medical School and then returned to Columbia to complete his PhD.

During his graduate years at Columbia, Muller would quickly become involved in impactful work with fruit flies (Drosophila) that would forever influence his life direction. He worked in the famous "Fly Room" with geneticist [Dr. Thomas Hunt "T.H." Morgan (born 1866)], who brought Muller into his lab based on the reputation of Muller's prior work. Muller and Morgan had mismatched temperaments, with Morgan's more laissez-faire approach clashing with Muller, who did not like others getting credit for his work.

Despite their differences, Muller played an important role in the [Dr. Thomas Hunt "T.H." Morgan (born 1866)] laboratory. He wrote a series of papers on the mechanism of crossing-over of genes, and his thesis on crossing-over earning him his Ph.D. in 1916. Crossing-over refers to an exchange of genes between a pair of homologous chromosomes, resulting in a mixture of parental characteristics. Muller was also co-author, with Morgan and two other students, on The Mechanics of Mendelian Heredity, a landmark book that linked Gregor Mendel’s laws of heredity with discoveries about chromosomes, the bodies in the cell nucleus that Morgan’s group had shown to contain genes.

Despite the accomplishments of the [Dr. Thomas Hunt "T.H." Morgan (born 1866)] lab and Muller’s desire to continue to work at Columbia, he would soon be leaving. Foundation Chair [Sir Julian Sorell Huxley (born 1887)] of the Biology Department at the new Rice Institute was impressed with the work coming from the lab. On recommendation from Morgan, Muller did post-doctoral work at the Rice Institute in 1916 through 1917. He would return to Columbia from 1918 to 1920 where he would serve as interim professor while Morgan was on sabbatical. Muller’s hope was that a more permanent faculty position would open up for him, but perhaps through various issues surrounding his strained relations with Morgan, the faculty position was not offered. Muller would return to Texas, but this time, to UT Austin.

UT YEARS

In 1920, Muller was hired as an Associate Professor by Dr. John T. Patterson, who believed in experimental zoology. Unlike his situation at Columbia where Muller had to put much of his own money towards lab supplies, Patterson made sure Muller had money for equipment and an assistant. The new Biological Laboratories (BIO) building that opened in 1925 provided Muller the luxury of a refrigeration unit to make studying of flies much easier than it had been in his earlier building in the Old Main, as the Texas heat would sterilize the flies.

Muller had a profound effect on his colleagues at UT, specifically Patterson who was studying armadillo embryology and Theophilus S. Painter who was studying the cytology of spiders. When Muller showed them the use of Drosophila in genetic analysis, both professors switched their organisms of choice to this insect. While this increased the discussions and approaches, Muller soon felt he was putting too much time into his colleagues’ work and not enough into his own. In addition to his day hours, he would begin to work at night, seven days a week, sometimes sleeping on a cot. His contemporaries noticed Muller's avoidance of them, and this increased rivalry.

Muller's most important discoveries occurred between 1918 and 1926. With [Dr. Thomas Hunt "T.H." Morgan (born 1866)], he became interested in mutations, for these occurred so rarely in nature that they were hard to study. Muller also believed that mutations were mostly recessive and would appear when two organisms carrying the mutated gene mated and passed on that gene to their offspring. This might not occur until generations after the mutation initially arose. Mutations also might not become visible if they caused death during embryonic development.

Looking for a way to increase the frequency with which mutations occurred, Muller first tried heat in 1919 but found in 1926 that X-rays were far more effective. The machine that facilitated these X-ray experiments is still visible on campus, in one of the entrances of the MBB building.

In a ground-breaking paper published in 1927, Muller announced that exposure to X-rays increased the frequency of mutations in fruit flies to 100 times their natural level. This was the first time a large number of mutations had been produced artificially and would provide the potential to advance the study of genetics considerably. The artificial creation of mutations helped to lay the foundation for genetic engineering. Muller would be awarded the Nobel Prize in physiology or medicine in 1946 for this work.

In the second part to this article on Muller, we will take a look at the troubled times Muller experienced at UT before heading off to Europe.

SOURCES

• Carlson, Elof Axel. Genes, Radiation, and Society: The Life and Work of H.J. Muller. Cornell University Press, 1981.
• Carlson, Elof Axel. “Herman Joseph Muller 1890-1967. A Biographical Memoir.” National Academy of Sciences, 2009.
• Carlson, Elof Axel. “Speaking Out About the Social Implications of Science: The Uneven Legacy of H.J. Muller.” January 2011. Genetics Society of America.
• Crow, James F. Comment page in Discovery, 1994. Vol 13. No.4.
• Goodstein, Judith. “The Thomas Hunt Morgan Era in Biology.” Engineering & Science, Summer 1991.
• Green, Tim. “Hermann Muller: A genetics pioneer.” January 19, 2010. UTNews.
• Yont, Lisa. Facts on File Science Library: Notable Scientists A to Z of Biologists, Facts on File 2003.
• Wager, Robert P. and Crow, James F. “The Other Fly Room: J.T. Patterson and Texas Genetics.” Genetics Society of America, 2001.
• “Eighty years ago, Texas researcher conducted X-ray experiments that won Nobel prize.” September 28, 2007. UTNews.

TROUBLE IN TEXAS

As mentioned in Part One of this article, Hermann Muller was becoming a superstar in the science world of the 1920s. His talks and research on X-rays causing mutations in the fruit fly Drosophila were receiving world-wide attention. The size of his lab at UT Austin was increasing. But his long work hours and animosity towards his colleagues at UT were beginning to cause trouble. In addition, the development of politics and world affairs in the 1930s caused Muller to become attracted to communist-leaning ideals. This spelled trouble in a USA very much opposed to these ideas.

Muller had met Jessie Jacobs, an instructor in pure mathematics, while at UT. They would marry, and when Jacobs became pregnant with their first child, she would lose her position because at the time, her colleagues felt that a mother could not give full attention to classroom duties and remain a good parent. Jessie’s loss of her connections to her academic career would bring strain into the marriage, in addition to Muller's long night hours at his lab.

Muller’s relations with Painter and Patterson also began to deteriorate in 1929, with Painter skeptical of radiation causing reverse mutations, and Muller feeling Painter and Patterson were using his ideas and taking too much of his time. In addition, his formal mentor, Morgan, was still being acknowledged as the power center of genetics, despite all of the work Muller had done in Texas.

With the start of the Great Depression in the US, and the huge rise in unemployment, Muller felt burdened by the economic reality of many Americans suffering through this hardship. While most of his beliefs seem quite reasonable today (equality in wages for women, civil rights of African Americans), Muller's concern for the state of poor Americans was considered communist at the time. Even with the knowledge of the trouble this could possibly bring him, Muller served as faculty advisor of the National Student League, a student-led organization at UT that was also a communist front, advocating for things such as lower tuition rates, political and social equalities for minorities and blacks, and unemployment insurance.

On January 10, 1932, Muller wrote a note to fellow geneticist Edgar Altenburg. In the letter, Muller commented on his work as a scientist, writing that he was "too psychologically old to continue the struggle." He also requested that Altenburg give $1000 of Muller's money in support of the Communist Party. Muller then headed out to the woods outside of Austin and swallowed a roll of sleeping pills. He failed to show up to class the next day, and his wife frantically called his lab as he had not come home the night before. Several of Muller’s students and staff formed a search party, eventually locating him dazed under a tree but still alive.

While gossip about his suicide attempt spread, Muller quickly recovered, and in a complete turn of events, he won a Guggenheim Fellowship to study at the Institute of Brain Research in Berlin. This was very much a relief, for Muller could sense his days in Texas were numbered.

In his paper "The Dominance of Economics over Eugenics," Muller had quite publicly condemned the eugenics movement at the Sixth International Congress of Eugenics, arguing against current eugenics assumption that socially undesirable traits like vagrancy were innate. Muller instead argued that socially undesirable traits were a result of social situations. At the time, this was a very Marxist position, and considered outrageous.

In addition, Muller had by now opted to sponsor the newspaper the UT National Student League wanted to distribute. It was called "The Spark" and represented the left-wing views of the League, but it would not be permitted at the university. Despite the possible repercussions he could face at the time, Muller opted to help get it into print and distributed. He was unaware that the FBI had had infiltrated the National Student League, and also had Muller under surveillance. The FBI made this involvement known to then President of UT Austin, who was only relieved to know Muller would be leaving soon for Berlin.

With all of this stress, by the end of 1932, Muller and his wife, Jessie, had agreed to a separation.

GONE FROM TEXAS

With these mounting pressures, Muller went overseas to Germany. However little he took Hitler seriously at first, Muller soon faced the reality of the danger of Nazi takeover when the Nazis vandalized the institute where Muller was working. Muller opted to accept an invitation to establish a genetics laboratory in the Soviet Union where interest in genetics was growing.

Muller's five years in the Soviet Union changed a lot for him. Free of teaching duties, he became senior geneticist at the Soviet Academy of Sciences’ Institute of Genetics in Leningrad and Moscow. He had a lot of support and built a lab to initiate a focus on gene function explored through the "position effect," the position effect implying that the functioning of a gene is to a certain extent at least dependent upon what other genes are in proximity. Muller felt an admiration of Soviet life and socialism, and even Jessie and their son David came to live in an attempt to reconcile the marriage.

Despite this upturn in events, trouble would find Muller again. The move to the Soviet Union did not help Muller's marriage, and it was dissolved in 1935 after Jessie and their son returned to the States.

Muller found more trouble in his attempts to launch a eugenics program in the Soviet Union. Muller wrote a letter to Stalin advocating his ideas of a positive eugenics program, but there was a counter movement occurring. Trofim D. Lysenko offered a different view of heredity, based on his experiments with plant breeding, and upheld Lamarckian ideas in which the environment modifies genetic material in a direct or predictable way. Lysenko viewed western genetics, and thus Muller's views, as capitalist, racist, and bourgeois.

A very heated debate would occur, with Muller on one side and the Lysenkoists on the other. Simultaneously, many of Muller's colleagues would suffer from Stalin's purges, accused of being Trotskyites, leading to their arrests and execution. In a mass meeting of 3000 people, Muller debated Lysenko, essentially calling him a fraud, and causing an uproar amongst those supporting the Party agenda, and thus Lysenko. Advised to leave for his own safety, Muller enlisted as a volunteer in the Spanish Civil War in 1937 for the International Brigade, to combat Franco's fascist rule. Here he would do physiological research on blood transfusion.

Staying in Spain through the siege of Madrid, Muller needed to find a new place to go but had few options. His involvement in the communist-leaning UT student paper would force him to stand trial if he returned to UT. He could not return to the Soviet Union or face the possibility of execution. His wish was to work in Paris with Joliot Curie or in Stockholm with Gunnar Lundburg but there were no job openings.

Through the influence of [Sir Julian Sorell Huxley (born 1887)], in 1937 Muller joined the Institute of Animal Genetics at the University of Edinburgh, Scotland, where he developed a graduate program. Muller began to research the relationship of radiation dose to mutation frequency. He would advocate radiation safety and human use at a time when it was largely misunderstood. Muller remarried in Scotland and earned a doctorate of science in genetics in 1940.

However, Europe's tense midcentury political landscape would drive him once again from his home. World War II broke out, and Great Britain essentially shut down research in an effort to focus on the war at hand. In 1941, Muller moved to Amherst College in the US. During the war's slimming effect on research, Muller consulted on radiation projects for the Manhattan Project, in addition to teaching undergrads, something he did not care for much.

At the end of the war, with the knowledge that he would not be added to the Amherst faculty, he appealed desperately to friends to pull strings. Luckily, his reputation opened the door to become faculty at Indiana University in Bloomington, Indiana.

Muller would spend the rest of his teaching years here, and these were some of his happiest. He felt appreciated by his colleagues and was awarded the Nobel Prize in Physiology and Medicine in 1946, which had a transforming effect on his career. He was able to work on new problems in human genetics. He also had a daughter with his second wife.

During his tenure at Indiana University, Muller spoke out against radiation abuses during a time when this stance was read as a political attack against national security, and thus defending Communism. In an effort to protect his students and colleagues, he also burned much of his correspondence with communists during this era of McCarthyism, and had to testify before the House of Un-American Activities.

Muller also revived his ideals of eugenics, and felt that it could be used essentially like family planning to produce healthier, more intelligent and caring future generations. But in the shadow of the Holocaust and early American eugenics movements, his beliefs on how to apply this were very often misunderstood and as a result, condemned.

In 1964, Muller would suffer a severe heart attack that affected his health greatly. In the following years, his health would worsen, eventually leading him to the hospital for kidney failure on his 76th birthday. Muller’s condition did not improve, and he decided that he no longer wished to live, something he accomplished by refusing to eat. He died in Bloomington on April 5, 1967.

Herman Muller was a scientist passionate about his work, and equally passionate about speaking out against the abuses of science. He helped usher the careers of many of his students on whom he made a deep impression. Muller also found himself directly involved in some of the most tumultuous times in history, from the era of the Great Depression and tolerance of racism in the US, to the anti-Semitism and book burnings of Nazi Germany, to Stalin's purges. Despite his troubled and intense life, Muller left a lasting impression on the field of genetics, much of this influential work happening while here on the UT campus.

INTERESTING TRIVIA

• - UT’s first female Ph.D. in Zoology (1931), Bessie League, assisted in Muller's lab. Her office number in BIO was room 313.

• - Muller and science fiction writer Ursula Le Guin are second cousins.

• - Carl Sagan worked in Muller’s lab at Indiana University during the summer of 1954.

• - At a time when air conditioning was decades away, the "constant temperature room" in UT’s "Fly Room" was kept at 69 degrees year round for raising Drosophila during the hot Austin summers.

• - Muller’s granddaughter, Chandra Muller, is a professor in the Department of Sociology at UT.

• - Muller’s son, David E. Muller (1924-2008), speaks frankly about his father in these video recordings on the Oral History Collection project from The Cold Spring Harbor Laboratory (http://library.cshl.edu/oralhistory/interview/scientific-experience/becoming-scientist/discussions-his-father/)

SOURCES

• Carlson, Elof Axel. Genes, Radiation, and Society: The Life and Work of H.J. Muller. Cornell University Press, 1981.
• Carlson, Elof Axel. “Herman Joseph Muller 1890-1967. A Biographical Memoir.” National Academy of Sciences, 2009.
• Carlson, Elof Axel. “Speaking Out About the Social Implications of Science: The Uneven Legacy of H.J. Muller.” January 2011. Genetics Society of America.
• Crow, James F. Comment page in Discovery, 1994. Vol 13. No.4.
• Goodstein, Judith. “The Thomas Hunt Morgan Era in Biology.” Engineering & Science, Summer 1991.
• Green, Tim. “Hermann Muller: A genetics pioneer.” January 19, 2010. UTNews
• Yont, Lisa. Facts on File Science Library: Notable Scientists A to Z of Biologists, Facts on File 2003
• Wager, Robert P. and Crow, James F. “The Other Fly Room: J.T. Patterson and Texas Genetics.” Genetics Society of America, 2001.
• “Eighty years ago, Texas researcher conducted X-ray experiments that won Nobel prize.” September 28, 2007. UTNews.

Many believe it’s because of the historic association of nuclear plants with nuclear weapons. But during the first two decades of nuclear power, people were more excited than afraid of it.

In his magisterial new book, Energy: A Human History, the Pulitzer-winning historian, Richard Rhodes, quotes the inventor of the first peaceful nuclear power plant, U.S. Navy Admiral Hyman Rickover, trying to tamp down excess enthusiasm by the public and policymakers.

“I think we have babied a lot of people in this country too long with the glamour of atomic energy,” Rickover told a congressman in 1957.

How did we go from the glamour of nuclear power in the 1950s to the fears that surround the technology today?

The most common answer to this question is that the nuclear accidents at Three Mile Island, Chernobyl and Fukushima frightened people.

And yet the accidents proved the relative safety, not relative danger, of nuclear energy. Nobody died from radiation at Three Mile Island or Fukushima, and fewer than 50 died died from Chernobyl in the 30 years since the accident.

How, then, did everyone come to see those nuclear accidents as so catastrophic?

The answer is because of how governments responded to them. Instead of encouraging the public to stay calm and carry on, governments freaked out, and evacuated hundreds of thousands of people.

“Between five and ten times too many people were moved away from the Chernobyl area between 1986 and 1990,” a team of top scientists wrote last week in the peer-reviewed journal, Process Safety and Environmental Protection.

As for the 2011 nuclear accident in Japan, the scientists said they found “it difficult to justify moving anyone away from Fukushima Daiichi on grounds of radiological protection.”

In other words, it was the over-reaction to the accidents — not the accidents themselves — that resulted in popular fears of the technology.

The statement by radiation scientists raises the possibility that, even if we cannot ever fully overcome public fears, we might change how governments respond to the next nuclear accident.

But it also begs the question: why do governments keep overreacting to nuclear accidents in the first place?

To answer that question, we have to go back in time to the birth of nuclear power — and the 50 year-long war against it.

The War on Universal Prosperity

In his 1953 “Atoms for Peace” speech, President Eisenhower proposed using nuclear energy as a way to redeem humankind for having brought such an awful technology into existence. Arms reduction wouldn’t be sufficient. What was the point of peace if billions remained in poverty?

“Experts would be mobilized to apply atomic energy to the needs of agriculture, medicine and other peaceful activities,” Eisenhower told the United Nations General Assembly in Manhattan, New York. “A special purpose would be to provide abundant electrical energy in the power-starved areas of the world.”

Eisenhower’s vision was at once nationalist and internationalist, altruistic and self-interested.

“The United States pledges to devote its entire heart and mind,” he said, "to find the way by which the miraculous inventiveness of man shall not be dedicated to his death, but consecrated to his life.”

After Eisenhower finished, there was a brief silence and then something extraordinary happened: representatives from every nation rose to their feet and applauded for 10 minutes.

But not everybody was enchanted by the idea of eliminating poverty. Three years before Eisenhower’s speech, a veteran of the Manhattan Project, the U.S. government effort to create the atom bomb, published a book that argued that humans were overpopulating the earth.

Humankind “would not rest content until the earth is covered completely, and to a considerable depth, with a writhing mass of human beings, much as a dead cow is covered with a pulsating mass of maggots,” the scientist Harrison Brown wrote in The Challenge of Man’s Future in 1950.

Brown, notes Rhodes, was hugely influential among environmentalists. One of his protégés was John Holdren, President Barack Obama’s science advisor, who would go on to describe Brown as “warm and witty…and surprisingly modest.” But Brown had also proposed the breeding and sterilization of humans to prevent “the long-range degeneration of the human stock.”

Brown’s proposal, Rhodes explains, was an extension of the ideas of 19th Century economist Thomas Malthus who lusted for the extermination of his fellow man, particularly the poor and the Irish. “Instead of recommending cleanliness to the poor,” Malthus argued, “we should encourage contrary habits…and court the return of the plague.”

In 1966, misanthropic conservationists within the Sierra Club had embraced Malthusianism. Writes Rhodes:

The small-world, zero-population-growth, soft-energy-path faction of the environmental movement that emerge across the 1960s and 1970s knowingly or unknowingly incorporated the antihumanist ideology of the neo-Malthusians into its arguments… “more power plants create more industry,” [the Sierra Club’s executive director complained,] “that in turn invites greater population density.”

Such anti-humanist ideas came full bloom in Stanford biologist Paul Ehrlich’s 1967 Sierra Club pamphlet, The Population Bomb, which depicted poor people in India as animals “screaming…begging…defecating and urinating.”

In sharp contrast, the creators of nuclear power remained optimistic and humanistic. They viewed the new energy source as the key to avoiding the problems created by a growing human population — and allowing everyone, including the poorest of the poor in Africa, to rise from poverty.

With nuclear energy, Oak Ridge Laboratory Director, Alvin Weinberg, argued, humans could create fertilizer, fresh water, and thus abundant food — forever.

But literal-minded nuclear advocates like Weinberg missed the point. Cheap and abundant energy was — for Malthusians — not a feature but rather a bug. The Sierra Club and other environmentalists hated nuclear because it held out the promise of universal prosperity.

It was at that moment that environmental groups and their philanthropic supporters began a half-century long campaign to frighten the public [see https://environmentalprogress.org/the-war-on-nuclear/]. "Our campaign stressing the hazards of nuclear power,” wrote Sierra Club’s President [see https://environmentalprogress.org/big-news/2017/3/28/why-the-war-on-nuclear-threatens-us-all ] in a 1974 memo to the board of directors, “will supply a rationale for increasing regulation and add to the cost of the industry."

The Scientific Fraud

One of the most shocking passages in [Richard] Rhodes’ Energy [see https://www.amazon.com/Energy-Human-History-Richard-Rhodes-ebook/dp/B075RVX2N3] is about how a Nobel Prize-winning American scientist committed scientific fraud to exaggerate the risks of nuclear radiation to human health.

Drawing on archival research by Edward Calabrese, a University of Massachusetts, Amherst professor of toxicology, Rhodes describes the work of Hermann Muller, a University of Texas geneticist who won the Nobel Prize in Medicine in 1946.

Muller’s research on fruit flies led him to conclude that there is no safe dose of radiation because every dose, he believed, led to mutations that, Rhodes explains, “were damaging or lethal… irreversible and permanent as well.”

But just before Muller flew to Stockholm to accept his Nobel Prize, he was sent new research that contradicted his conclusions.

Muller’s work, and that of others across the years, had explored the effects of high and medium doses of radiation. [Insect behaviorist Ernst] Caspari had extended that research into the low-dose range and had asked in particular whether the effect would be the same when the dose was spread out over a period of time (“chronic”) rather than delivered all at once (“acute”)...Caspari’s startling new finding was that fruit flies exposed to a [low] daily dose… showed no increase in their mutation rate.

Muller faced a dilemma. “What should he do? What he should have done was qualify his Nobel lecture.” But Muller didn’t. “In Stockholm,” Rhodes writes, “Muller accepted his Nobel Prize and then deliberately ignored Caspari’s findings in his lecture.”

But that wasn’t even the worst of it. On his return to the U.S., Muller reviewed Caspari’s research and told a colleague that he had “little to suggest in regard to the manuscript” beyond recommending the study be replicated.

However, as the main reviewer of the paper, Muller proceeded to oversee its publication — with two changes. “Muller’s name now appeared among the acknowledgments,” notes Rhodes, “and one crucial sentence had been deleted. The deleted sentence was the sentence that questioned Muller’s theory.”

Muller’s status as a powerful scientist and Nobel Prize winner allowed him to establish his falsified theory as as the scientific basis for regulating nuclear plants for decades to come.

Having suppressed an evidence-based challenge to his “linear no-threshold” (LNT) model of radiation effects, Nobel laureate Muller thereafter continued to promote and defend the LNT model whenever and wherever the question arose.

Muller’s professional motivations overlapped with the agendas of anti-nuclear scientists and activists. “An antinuclear movement that originated in hostility to population growth in a supposedly Malthusian world,” writes Rhodes, “promoted in turn the LNT model, exaggerating its effects.”

The good news is that a growing number of scientists who specialize in radiation, climate, and public health are speaking out for nuclear power plants as critical to saving lives.

In 2013, Rhodes notes, the climate scientists Pushker Kharecha and James Hansen found that “nuclear power prevented an average of 1.84 million air pollution-related deaths.”

And that’s before taking into account the potential consequences of climate change.

Over the last two and half years, climate scientists like Hansen and scholars like Rhodes have joined forces to protect and expand nuclear power plants, from Illinois and New York to South Korea and France.

Now, radiation scientists with the backing of the British and Indian governments are urging governments to stay calm and carry on during nuclear accidents.

Their efforts hold out the hope that, whether or not fears of air pollution and global warming can ever trump fears of nuclear accidents, we might at least stop ourselves from grossly over-reacting to them.

Early Life

[ Source pages (saved as PDF) - [HE084][GDrive] / [HE088][GDrive] ]

Muller was born to a working class family in New York City in 1890.

His father, Hermann Sr., had taken over the family metalworks shop abandoning a nascent career in law, when his own father, a German immigrant, died.

Hermann Sr. had strong interests in science, socialism, and emancipatory politics. Despite his unexpected death in 1900 when his son was only 9 years old, he was able to pass these interests onto his young son. Hermann considered his father instrumental in inspiring him to be a scientist, as he wrote in his autobiographical notes.

Growing up, he embraced a wide array of scientific interests from bug collecting to stargazing. Newspaper clippings in his childhood journal, indicate the breadth of his early scientific and socio-political interests. Meanwhile his mother Frances, of English and Sephardic Jewish descent, and a school teacher, encouraged Hermann with his studies.

Muller attended Morris High School in the Bronx, where he met his lifelong friend and future fellow geneticist Edgar Altenburg. They helped found the Morris Science Club. An excellent student, Muller was school valedictorian and spoke on "The Need for Higher Ideals in Business and Politics" at his graduation. Society's lack of higher ideals frustrated him throughout his life.

Supported by a city scholarship, Muller started at Columbia University in 1907 at age 16. He pursued his passions for politics, joining the Intercollegiate Socialist Society, and for science, studying biology. The Columbia biology department, chaired by the renowned zoologist, cell biologist, and early geneticist E.B. Wilson, was considered one of the best in the nation. The introductory biology class, taught by a former student of Wilson, J. H. McGregor, introduced Muller to the power of evolutionary theory--something that had been absent from his high school education. Wilson himself was a giant in Muller’s life. His own course covered a great deal of what had only recently begun to be called genetics.

New Science of Genetics

[ This section source (saved as PDF) - [HE008C][GDrive] ]

In the first decade of the twentieth century, biology was in a period of great upheaval. One of the mysteries that emerged in the aftermath of Darwin was heredity. If organisms had developed over millions of years by passing along their traits with slight modifications in a sequence of tiny steps that ultimately resulted in vastly different species, then how exactly were traits passed from parents to offspring? And how were novel traits not simply swamped out by being blended with other traits?

Just a handful of years before Muller attended Columbia, three separate scientists, Carl Correns, Erich Tschermak, and Hugo de Vries, had rediscovered the 1866 work of the Austrian monk and plant breeder Gregor Mendel. Mendel had found that inheritance may be “particulate,” or inherited in discrete units. This contrasted with prevailing theories of blended inheritance, in which traits in offspring are an intermediate mix of parental traits. Mendel observed for instance that if a yellow pea plant is crossed (mated) with a green pea plant, rather than producing greenish yellow peas, as would be expected with blended inheritance, one instead finds a fixed ratio of green pea plants to yellow pea plants in the offspring.

This tale of “rediscovery” is traditional in histories of genetics, but also a bit misleading. Far from simply stumbling across Mendel’s work in a rarely used part of the library and realizing its significance, the rediscovers were involved in well-developed experimental programs of their own and only slowly came to the realization their work aligned with Mendel’s.

Beyond observable patterns of inheritance, early twentieth century genetics was also interested in the ultimate physical means of inheritance. Wilson was one of the leading advocates of chromosome theory, the doctrine that the thread-like bodies which could be observed through microscopes multiplying and dividing during cell division, were in fact the carriers of the hereditary material. Wilson and his graduate students interpreted the separation and division of the chromosomes as physical causes behind Mendelian inheritance.

For Muller, Wilson’s undergraduate class was a thrilling experience. It covered the latest findings in biology and unified them under the aegis of chromosome theory. Muller became a true believer in chromosome theory and the new science of genetics. In a debate society he joined with his friend Altenburg, he also explored the political implications of modern biology, a topic that concerned him throughout his life. In fact, almost twenty years later his college debate talk "Revelations of biology and their significance" would be expanded into his book Out of the Night.

The "Fly Room" at Columbia

[ This section sourced from (and saved as PFD) - [HE008F][GDrive] / [HE008I][GDrive] / [HE008K][GDrive] ]

Muller graduated from Columbia University in 1910. Then, working part-time as a bank runner and an English teacher, he enrolled as a fellow in physiology at Columbia Medical College. He took a full load of courses, including those of the well-known embryologist and up-and-coming geneticist Thomas Hunt Morgan. Muller would never have the enthusiasm for Morgan he had for Wilson, but in those years of 1910-1912 he spent a great deal of the time in Morgan’s lab -- known as the “fly room” -- and became a formal member of the lab in 1912. Alongside [Dr. Alfred Henry Sturtevant (born 1891)] and Calvin Bridges, Muller counts among the early group of graduate students that helped catapult the lab to fame.

The Columbia "fly room" was so named because genetics was studied in the cramped space of 613 Schermerhorn Hall by breeding and experimenting on the tiny common fruit fly, Drosophila melanogaster.

Stories about [Dr. Thomas Hunt "T.H." Morgan (born 1866)]’s lab abound, but what is not debatable, is that the lab produced some of the most groundbreaking research in early twentieth-century genetics and launched fly genetics as a leading specialty within the field. For almost four decades, Drosophila melanogaster would be the dominant beast of animal genetics and a way of life for fly geneticists. Its importance continues up to the present.

Fernandus Payne, a graduate student who had recently graduated from a masters program at IU, introduced the Morgan lab to fruit flies around 1910. Payne becomes important again much later in this story, as he is the primary force behind IU hiring Muller.

Early on in his fly research, [Dr. Thomas Hunt "T.H." Morgan (born 1866)] found a fly that had white eyes rather than the standard red. Mutant strains like “white” were invaluable, as they enabled the presumed gene(s) behind the mutation to be tracked through crosses. Muller and the other fly room members became experts at spotting and breeding mutant fly strains. Together with the rise of fly genetics there was a proliferation of hundreds of highly cultivated mutant strains each with their own particular features suited to different experiments, as well as a national and international fly trade that facilitated experimental access to the right strains. Today IU Bloomington has one of the largest Drosophila stock centers in the world.

This draft of a paper on the fourth chromosome of Drosophila became Muller's first scientific publication. The patchwork penmanship stays with him for his whole career. It can even be found in his childhood writings.

Muller’s relationship with the fly room was complicated. First, he was simply busier than some of the other members of the lab--juggling several other jobs ensured he was unable to fully participate in laboratory culture. Unlike his colleagues Calvin Bridges and [Dr. Alfred Henry Sturtevant (born 1891)], Muller was not at first financially supported by [Dr. Thomas Hunt "T.H." Morgan (born 1866)]’s lab. More problematic were his intellectual issues with Morgan. In Muller’s view, Morgan’s saving grace was that he had the good sense to listen to his students. However, Morgan’s skepticism about chromosome theory and stubborn demands for straightforward experimental demonstration were endlessly irritating to the young Muller who had an uncanny gift for envisioning complex breeding experiments and what they said about chromosome structure. And it was no doubt difficult for Muller to give much credit to Morgan for adopting chromosome theory when E. B. Wilson, whom Muller greatly admired, had done so years earlier.

The one discovery Muller gave [Dr. Thomas Hunt "T.H." Morgan (born 1866)] indisputable credit for--and arguably the most important discovery to come out of the lab--is the phenomenon of crossing-over. The great insight of Mendelism was that inheritance was particulate, meaning that crosses would result in fixed ratios in offspring with distinct traits rather than just intermediate traits. One of the associated assumptions was that traits were inherited independently. For instance, if a pea plant had a ¼ chance of being yellow, and a ¼ chance of being wrinkled, then it had a 1/16 chance of being yellow and wrinkled. However, experimentalists like Thomas Hunt Morgan were finding that certain traits were inherited together far more often than would be expected by chance. It was this evidence, together with the urging of his students, that finally convinced Morgan to fully adopt chromosome theory. The fly room members reasoned that if genes for two separate traits were on the same chromosome, that would explain why they were usually inherited together.

Leaving the USSR

[ Source is (and content saved to PDFs as : [HE008L][GDrive] / [HE008Q][GDrive] / [HE008V][GDrive] / [HE008Z][GDrive] ]

Shortly after the debate, in early 1937, Muller heard that Stalin had read his book, Out of the Night. Stalin had rejected Muller’s vision of socialist genetics and was apparently readying an attack on the book. Having managed to personally irritate Stalin, and with no clear career prospects there, Vavilov urged Muller to leave the Soviet Union. However, he wanted to recoup some goodwill first to minimize the negative consequences of his departure to his colleagues. The plan was to join the Republicans in their fight against fascism in the Spanish Civil War, an international cause celebré.

The Soviet Union was the only major power that supported the rightfully elected government in Spain. The leftist Spanish Republicans were struggling to match the military aid provided by Hitler and Mussolini to the fascist insurgency led by Generalissimo Franco. Muller and Vavilov correctly felt they would not refuse volunteers. Muller joined the blood transfusion unit led by the renowned Canadian thoracic surgeon and political radical Dr. Norman Bethune, bringing with him Russian blood transfusion technology.

Canada, like the United States, France, England and many other nominally anti-Fascist powers had recognized the Republican government, but chose to sit back and watch rather than offer aid. However, there was a groundswell of support among leftists in these countries and many of them went on their own, to fight, to help, or to document history. Bethune himself would die a few years later working as a battlefield medic in revolutionary China. Back in the USSR, people were closely following the Spanish Civil War. Vavilov in fact wrote to Muller that his heroic rush into a collapsing building to help rescue some research had appeared in the Soviet papers.

Muller stayed in Spain for eight weeks and then briefly headed back to Texas, before returning to the Soviet Union. Having established his leftist bonafides with his time in Spain, Muller escaped after a final farewell with Vavilov in October 1937. Lysenkoism continued to grow and fester in Soviet genetics for the next several decades, being adopted as official doctrine in 1948, before finally being overcome in the 1960s, when Lysenko was removed from his position and Mendelian genetics could again be taught openly. The personal toll on Muller and his circle of friends and colleagues would be immense. Both of Muller’s former Soviet lab members from Texas, Izrail Agol and Solomon Levit, were shot on the “killing fields”, possibly before Muller had left. Muller’s friend and sponsor, Nicolai Vavilov, died of starvation in a Soviet prison in 1943 in the town of Saratov, near a university where he used to work. Many other geneticists shared their fate. Muller's experiences in the USSR closed the curtain on his communist sympathies, although he remained committed to broadly leftist politics for the rest of his life.

Given the sensitive nature of the political situation in the USSR, Muller was extremely quiet about the events that had occurred there. A careless word could have led to the death or imprisonment of a friend or colleague. Beyond the letter to [Sir Julian Sorell Huxley (born 1887)], Muller also provides a fairly detailed account in a 1944 letter to the geneticist Karl Sax, who was writing on the Lysenko controversy. The rose-colored glasses through which many fellow leftists viewed Lysenkoism and the political situation in the Soviet Union under Stalin, coupled with his inability to speak openly and correct their misconceptions would frustrate Muller for years to come.

One of his major research works from Soviet Russia, a bibliography of Drosophila research, Muller had to smuggle out of the country. The original is now held by the Lilly library. An edited version was later printed in Edinburgh in 1939 as the Bibliography on the Genetics of Drosophila. In a certain sad irony, the book was especially valued for its coverage of the excellent Russian work on Drosophila genetics.

Muller had no clear job lined up when he left Russia. He temporarily stayed in Paris with the Russian-born French geneticist Boris Ephrussi at the Radium Institute (now Curie Institute). [Sir Julian Sorell Huxley (born 1887)] in England meanwhile spoke with the English geneticist Francis Crew and got Muller an offer for a temporary position at Edinburgh.

Ephrussi would have liked to hire him as well and was eventually able to get together funds for an offer of his own, but Muller accepted the position at the Institute of Animal Genetics in Edinburgh. Crew, a pioneer in medical genetics, had been assembling a truly international institute, with researchers from India, Egypt, Germany and elsewhere. It was a good time to be in the recruiting business as many diligent and talented scientists were fleeing from fascist Europe. The Institute benefited greatly from members such as the German-Jewish Charlotte Auerbach and the Italian-Jewish Guido Pontecorvo.

Hitler was still ominously gaining power to the east, but that was no excuse to not focus on the business of genetics and Muller returned to normal work with enthusiasm. His student Sachi Prasad Ray-Chaudhuri was exploring the effects of radiation dose on mutation and Charlotte “Lotte” Auerbach engaged in her own trail-blazing research on the chemical induction of mutations with Muller’s encouragement and mentoring. She would become the first researcher to unequivocally establish that things other than radiation, in her case mustard gas, caused gene mutations. Of particular note is that Muller and his colleagues were increasingly finding evidence of the dangers of radiation, even for the small doses used in medical radiology. Despite their almost complete ignorance of the relevant science, many members of the medical community were highly recalcitrant and this would be a fight Muller would have to pursue over the next several decades.

Further distracting him from the looming threat of war, was a budding romance. Auerbach had introduced Muller to Dorothea “Thea” Kantorowicz, a German-Jewish refugee and physician then working as a pregnancy diagnostic technician at the same Institute. Theirs was a whirlwind romance and they married in May of 1939, less than a year after meeting. Thea deserves particular mention in this short biography, as besides her importance in Muller’s life, her efforts were also instrumental to the development and organization of the Muller collection at the Lilly Library where she voluntarily worked with his collection for ten years.

The Seventh International Genetics Congress that Vavilov had desperately hoped Muller could bring to the USSR followed Muller to Edinburgh instead. In a symbolic gesture, an empty chair on the podium was left for Vavilov, the elected chairman of the Congress, during the opening ceremony. The large Soviet delegation, planning to present some fifty papers, was forbidden to attend due to the continued politicization of genetics research in the Soviet Union

The Congress came with all the anxieties one would expect of an international gathering on the eve of a world war. Many papers were read in absentia, delegations had to leave suddenly, and the conference was concluded early. Several delegates sought to return home on the transatlantic British passenger liner SS Athenia, which, tragically, was confused for an armed vessel and torpedoed by a German U-boat hours after the declaration of war. There were 118 casualties, including an American geneticist and his wife.

In response to Nazi eugenics and other terrifying movements that claimed to make use of genetics, Muller drafted, and 22 other geneticists signed, the “Geneticist’s Manifesto.” The formal statement highlighted the relevance of genetics (and eugenics) to human well-being, but attacked the racist and dehumanizing tendencies of much of eugenics. Muller sent a copy of the manifesto to Vavilov. His response is the last of their communication.

Returning to America

[ Source (and saved to PDF as) : [HE0093][GDrive] ]

The tense situation in Edinburgh only intensified after the Congress, and Muller found it difficult to be a practicing scientist during WWII. As a foreigner, Muller’s movements were severely restricted, while Edinburgh seemed less safe by the day. Thea had to convalesce from tuberculosis in a hospital or sanitorium much of their time in the city and her “stateless” position made emigration problematic. The Mullers were informed at the last minute of a way out of the country, and the two of them planned an escape to New York City via London. However, the British port was bombed shortly before their departure and they had to abruptly divert to Lisbon and wait for the last Pan American seaplane to take them to the United States. Muller kept in correspondence with Francis Crew and other Edinburgh scientists throughout the war. Most of the men were drafted as soldiers or otherwise served in the war effort.

After a short stay in New York, one of the geneticists who had been on the torpedoed ship, Harold Plough, managed to get Muller a temporary position at Amherst College while Muller continued to search for a more permanent position. The job search was difficult and Muller was haunted by his past and concerns about his possible communist sympathies and even his (distant) Jewish ancestry. Scientists were generally ready to ignore inconvenient politics for scholarly acumen, but image-conscious administrators were not always so flexible. America’s entrance into the war left more room in the Amherst biology department for Muller and took some of the immediate pressure off the job search.

Unfortunately, the general absence of assistants and material (almost everything was being consumed by the war effort) and the teaching demands at an Amherst short on instructors slowed his research to a crawl. His time at Amherst was also a period of highs and lows in Muller’s family life. His sister, Ada, died in 1942, and Thea’s tuberculosis returned for a period in the early 40s. But his son David married, and Thea’s health improved enough to give birth to their daughter, Helen, in 1944.

The Indiana Years

[ Source (and saved to PDF as) : [HE0098][GDrive] / [HE009E][GDrive] / [HE009K][GDrive] / [HE009P][GDrive] / [HE009V][GDrive] ]

Just as the job prospects seemed hopeless, Fernandus Payne, the former [Dr. Thomas Hunt "T.H." Morgan (born 1866)] student and a longtime IU faculty member, pushed for the university to offer Muller a job. Payne, who was Dean of the University Graduate School and chair of the zoology department, had cultivated an excellent genetics program and knew they would benefit from Muller’s experience and expertise. IU’s president Herman B Wells was worried about Muller’s political past, but with Payne’s urging, ultimately embraced the opportunity to acquire such an eminent scientist. The papers of both Payne and Wells can be found in the Indiana University Archives.

Muller would stay at Indiana University for almost the rest of his career. He was far from the only established scientist in the program and his colleagues included the botanist Ralph Cleland, the microbiologist Tracy Sonneborn, the biologist-turned-sexologist Alfred Kinsey, and the future Nobel Prize winning bacteriologist Salvadore Luria. Muller arrived in Bloomington in the summer of 1945 and set up a sprawling Drosophila lab in Science Hall (now Lindley Hall).

IU could not have offered Muller a job at a better time. The very next year he was awarded the Nobel Prize for his research on X-rays and mutations. Muller had been out of town for the official telegram, so for several days he dealt with reporters calling him and asking for statements, and warm congratulations from colleagues, while still trying to confirm he had actually won it. In his Nobel speech he continued to emphasize his perennial theme of the importance of genetic understanding for the future of humanity.

The Prize was fortuitously timed for Muller as well. In the wake of the atomic bombings of Hiroshima and Nagasaki, radiation, atomic energy, and nuclear armaments became matters of international concern. Additionally, a particularly rabid form of anti-communism hatched in the post-war years as the Soviet Union emerged as America’s rival. The gravitas provided by the Nobel Prize would help Muller greatly in the years ahead as he used his scientific position to combat anti-communist witch hunts (Muller was subpoened to testify before the House Committee on Un-american Activities in 1953 where he was exonerated), the American government’s endless hunger for all things nuclear, and the international echoes of Lysenkoism.

In his address, at an IU convocation in his honor as Indiana University’s first Nobel Laureate, Muller warned about the dangers of radiation and what the power of the atomic bomb meant to civilization. Several drafts are in the collection. As a geneticist, he specifically worried about the effects the atomic bomb would have on not just those in the blast radius, but the more subtle effects of genetic damage. He stressed that mutations did not look like they did in the comics, or even in medical texts, but rather they could cause minor heritable harms whose full impact would be measured over hundreds or even thousands of years as a slow drag on humanity.

Pushing for the acknowledgement of the dangers of radiation as a scientist-activist is one of his most important achievements. He was opposed every step of the way. Physicians had been reluctant to acknowledge the harms of radiation and grew increasingly defensive, as Muller became increasingly accusatory. The federal government and the armed forces represented even more of an entrenched interest on the matter. Things came to a boiling point at the 1955 United Nations International Conference on the Peaceful Uses of Atomic Energy.

As the foremost expert in radiation genetics, Muller had originally been selected by the United States Atomic Energy Commission to give a paper on the genetic dangers of radiation. However, his paper was unexpectedly withdrawn. When he communicated with the Atomic Energy Commission, they lied and told him the UN had withdrawn his paper. In fact, it was a commissioner of the Atomic Energy Commission, the physical chemist Willard Libby, who was suspicious of Muller’s political past.

Whatever Libby’s rationale, the AEC’s suppression of Muller’s paper and the scapegoating of the UN back-fired. The controversy appeared on the front page of the Washington Post, and led to additional attention to the conference, the Atomic Energy Commission, and radiation by both scientists and the public. Muller would continuously advocate for responsible guidelines for radiation exposure for the rest of his life.

Muller increasingly became a public voice for science, above all arguing for a scientific rationalism, which held that science needed to be safe from interference and censorship, and its results and implications not willfully ignored. This entailed staunch criticism of Lysenkoism. But in his home country, it also meant striking a careful balance between advocating for free and open discourse, and not being too tolerant of intolerant ideologies--among which he now included communism. Scarred by his time in the Soviet Union and the brutality of Stalin, as well as the continued international communist support of Lysenkoism, Muller had little sympathy left for the communist party. He was similarly distrustful of the Soviet Union in international politics, and while he was anti-war and often joined in anti-war causes with other eminent scientists, he thought disarmament without a concordant agreement from the USSR would be a mistake. He signed the 1955 Einstein - Russell Manifesto authored by Albert Einstein and Bertrand Russell that warned of the dangers of atomic weapons, but with qualifications. From 1956 to 1958 he served as president of the American Humanist Association, continuing to push his compassionate scientific rationalism. Muller was elected Humanist of the Year in 1963. Deeply entwined with his humanism, he also invested time in promoting germinal choice (sperm banking) as a way for humanity to have more control over reproduction.

Despite Muller’s rising status as an international public scientist and humanist from his Nobel Prize onward, in the trenches of genetics, classical genetics and Drosophila studies were losing pride of place. Beginning in the 1940s, new model organisms, like the fungus Neurospora, bacteria, and bacteriophages (viruses that infect bacteria), were becoming of increasing interest. The discovery of the structure of DNA in 1953 led to new excitement about studying the raw material of heredity directly, rather than mediated through breeding experiments. Despite the shift away from fly genetics, Muller was no member of the old guard. He had long been advocating for the integration of physics, chemistry, and genetics which heralded the new era of molecular genetics and encouraged the department to move in that direction. One of the discoverers of DNA, James Watson, specifically went to IU in 1947 because of Muller, where he took Muller’s advanced genetics course.

Muller’s work on the gene as the basis of life was the intellectual ancestor of the new exuberance about DNA, though not every molecular geneticist was aware of the full extent of this heritage.

Muller formally retired from IU in 1963, having done a great deal for the university, and a great deal more for genetics. He then took visiting scholar positions at the City of Hope in California and The University of Wisconsin. Muller continued to lecture, write, and occasionally teach until 1966.

Muller died of complications of heart disease at the Indiana University Medical Center In Indianapolis in April of 1967. There is no doubt Muller would be unhappy with the present state of the world, and especially America, with its rampant economic inequality and science denial. It is however a world that knows the dangers of radiation, rarely tests nuclear weapons, and understands the tremendous importance of genetics. It is also a world in which his children and students went on to build successful lives and careers. Perhaps then, even if he would be dissatisfied with the state of the world, Muller would be proud of the enduring impact he had on it.

Muller's paraphrase of Omar Khayyam (Possibly drawn from Edward FitzGerald's Rubaiyat of Omar Khayyam).

A glowing cloud formed a planet,

Then a gene, and life began--

At length, a cell, then a polyp,

A four footed beast, and a man:

Man by mutual learning and striving

Wins to far-reaching vision and plan,

Till at last he'll stride heedfully

Forward with paces of cosmic span.

n a t i o n a l a c a d e m y o f s c i e n c e s

h e r m a n n j o s e p h m u l l e r

1 8 9 0 — 1 9 6 7

A Biographical Memoir by

e l o f a x e l c a r l s o n

Any opinions expressed in this memoir are those of the author

and do not necessarily reflect the views of the

National Academy of Sciences.

Biographical Memoir

Copyright 2009

national academy of sciences

washington, d.c.

HERMANN JOSEPH MULLER

December 20, 1890–April 7, 1967

BY ELOF AXEL CARLSON

Hermann joseph muller is best known as the founder of the field of radiation genetics, for which he received the Nobel Prize in Physiology or Medicine in 1946. He was also a cofounder—with Thomas Hunt Morgan, Calvin Blackman Bridges, and Alfred Henry Sturtevant—of the American school of classical genetics, whose use of the fruit fly, Drosophila melanogaster, provided a remarkable series of discoveries leading to an American domination of the new field of genetics first named in 1906 by William Bateson.1 Muller’s career as a geneticist was productive and included 370 publications and participation in active laboratories in Texas (the Rice Institute and later the University of Texas at Austin), the Soviet Union (at their National Academy of Sciences in Moscow and Leningrad, Muller being a corresponding member), Edinburgh (the Institute for Animal Genetics at the University of Edinburgh), and Bloomington, Indiana (in the Zoology Department at Indiana University).2

Muller was a controversial critic of society who made an effort to decry abuses of genetics and who served on many national and international committees as an advocate for radiation safety. He was both a critic and advocate of eugenics, denouncing the American eugenics movement for its racism, spurious elitism, sexism, and mistaken assumptions on both the transmission of behavioral traits and the belief that many social traits were primarily innate. He promoted an idealistic eugenics throughout his life, believing that those with beneficial genes should have opportunities to transmit them. His early enthusiasm for socialism and communism cost him dearly later in life, despite his role as the leading world critic of the Lamarckian movement initiated in the Soviet Union by Trofim D. Lysenko.

FAMILY AND EDUCATION

Muller was born in New York City on December 20, 1890. His friends knew him as Herman (the last n being dropped) until the 1940s, when he shifted to Joe on the recommendation of his second wife, Dorothea (née Kantorowicz) Muller. Professionally, he used his initials and his articles appear as H. J. Muller. Muller was a third-generation American. His father’s ancestors came to the United States from Coblenz (the Rhine Valley) in Germany after the unsuccessful revolution of 1848, which they supported. Three Muller brothers came to the United States that year. One died the following year when he tried to make his fortune in the California gold rush. The other two brothers established themselves in an art metal business in New York City on Canal Street, preparing bas reliefs, frames, and other objects for the middle class homes of that era.

The Mullers were originally Catholic, but on H. J. Muller’s side they became Unitarians, the religion of upbringing of H. J. Muller. They were also sympathetic to the emerging labor and socialist movements, an influence that carried over to the young Muller. On his mother’s side (the Lyons) the family was originally from England of mixed Jewish and Anglican background. This less-than-half “Jewishness” Muller used on occasion to offer his solidarity with Jewish colleagues who were victims of anti-Semitism in the United States or Germany. Muller himself was an atheist and later in life chose the American Humanist Association as an outlet for his religious feelings, serving as president of that organization in 1957.

On his uncle’s side Muller’s penchant for academic life was shared by his first cousin Herbert J. Muller (English critic and author of Uses of the Past) and first cousin Alfred Kroeber (anthropologist known for his studies of American Indian cultures). Alfred Kroeber’s daughter became the wellknown science fiction writer, Ursula LeGuin. Muller had two children. With his first wife, Jessie (née Jacobs) Muller, he had a son, David Muller, who became a professor of mathematics. With his second wife (she called herself Thea) he had a daughter, Helen Muller, who became a professor of marketing and public health. David Muller has continued the academic tradition with a son, Kenneth Muller, a professor of neurobiology. Helen has also continued the tradition with a daughter, Mala Htun, who is a professor of anthropology.3

The young Muller attended Morris High School in the Bronx while commuting from upper Yorkville in Manhattan. He excelled in school and received a Cooper-Hewitt scholarship to attend Columbia University. His father died when young Muller was 10 years old, and the family lived on a modest income from the partnership with his father’s brother. During his college years, young Muller had to work odd jobs part-time to help support his mother and sister. At Columbia, Muller received his B.A. in 1911 and Ph.D. in 1916.

He knew he wanted to be a scientist and had considered engineering and basic science as possibilities when he entered Columbia. He quickly chose the life sciences after taking courses with Edmund Beecher Wilson. Morgan at the time had not yet established himself as a geneticist and Muller’s only course with Morgan did not mention his work on fruit flies.4 Muller felt intellectually excited by Wilson’s ideas on the cell and the chromosomes. He felt Morgan was muddled in his thinking on evolution and genetics. This surprising evaluation makes more sense to those who know that at the time when Muller was an undergraduate, Morgan was strongly influenced by the ideas and work of Hugo DeVries, one of Mendel’s rediscoverers and the proponent of the mutation theory.5 This theory believed new species arose de novo rather than by a gradual Darwinian change over hundreds or thousands of generations. Muller was a committed Darwinian and strongly supported natural selection as the basic mechanism of evolution, a view under academic attack in the early years of the 20th century.

While Muller was completing his bachelor’s degree, he met Bridges and Sturtevant, who had a very different experience with Morgan. Bridges, an orphan, was benefited by a part-time job as a bottle washer and food preparer for an organism Morgan had been studying for two or three years on recommendation of William Ernest Castle at Harvard. Morgan was hoping to find new species of fruit flies just as DeVries found new species in the evening primrose, Oenothera lamarckiana. Sturtevant, who lived with his brother’s family (he was a professor of linguistics at Columbia), impressed Morgan with his brilliance in class and with his initiative to write a paper on coat color inheritance in horses. Both Bridges and Sturtevant discussed Morgan’s recent finding of a mutation (white eyes) and its unusual mode of inheritance. Muller, who shared an enthusiasm for biology through the university’s biology club, was eager to join Morgan’s laboratory after his graduation (he was working on a master’s degree in nerve physiology at Cornell). Despite Muller’s personality, Morgan took him on. Muller had a reputation for his own brilliance, especially in coming up with powerful interpretations of the new findings from Morgan’s laboratory.

While a sibling rivalry simmered among the three graduate students, and Muller often was shunted to another room (to work with his lifelong friend and fellow high school alumnus, Edgar Altenburg, who was not accepted into Morgan’s laboratory), these budding geneticists engaged in numerous debates and discussions of all their experimental work.6 This makes the source of the ideas for their experiments and interpretations of the experiments difficult, if not impossible, to separate. It led to many disputes over priorities in their later careers, and a mutual hurt, each thinking the other greedy or unkind. Muller lamented that so much of his time that should have gone into carrying out the experiments of his ideas had to wait or was taken over by his rivals, because he was not supported by Morgan and he had to teach English to immigrants or work as a runner in Wall Street to earn money for his own and his mother’s needs. Sturtevant did not hesitate to lament that Muller had a “priority complex” and did not credit others for their own insights. Morgan sided with Sturtevant and Bridges; he took the view that ideas were cheap and commonplace and of little importance without the experiments to test them. It was Morgan who chose his own title, professor of experimental zoology. Morgan never felt embarrassed with his wrong ideas (he had many) because they fell by the wayside when he put them to test.

Although Muller finished his dissertation work in 1915, his degree was not awarded until 1916. His dissertation was on crossing over, a phenomenon first discovered in England and misinterpreted by Bateson as coupling and repulsion.7 Bateson thought of Mendelian units almost like bipolar magnets that repelled or attracted each other. A different model impressed Morgan, one that stemmed from his reading of a paper by F. A. Janssens on meiosis, which showed chromosomes twisted around each other. Morgan speculated that these twisted threads could break and reunite, separating or bringing together segments of a paternal and maternal homologue. He called the process “crossing over.”

Sturtevant used the data from Morgan’s first X-linked mutations and constructed a map. Muller was in awe of Sturtevant’s interpretation. (Sturtevant was still an undergraduate when he created the first map). Muller suggested using the ratio of crossovers to the total of crossovers and noncrossovers for determining the map unit. (Sturtevant had used the ratio of crossovers to noncrossovers). Muller then followed up the mapping of the mutations and determined that genes kept in heterozygous state for several generations did not lose their specificity. He also worked out a mechanism and interpretation, measured by what he called the coincidence and interference of crossing over for inconsistencies in map length. This resolved the observation that certain linked genes (those relatively close to one another) had a predictable distance when their individual internal distances were added, but those relatively far apart fell short of that predicted distance. In Muller’s interpretation, genes close together rarely had intervening double crossovers and those farther apart usually did. This made the longer distances shorter than the sum of the distances of contiguous segments within that stretch between the two outer genes. Muller believed the first break led to a release of tension nearby and this prevented multiple crossovers in that region.

GENETICS AT RICE UNI VERSITY

Muller decided to leave Morgan’s laboratory after a visit by Julian Huxley, who had been appointed as the founding chair of the Biology Department at the new Rice Institute. Huxley was impressed by the work of the Morgan school, and he asked Morgan to recommend a student; he recommended Muller for the job. Muller established a lifelong friendship with Huxley in those few years they had together at Houston. It made each a strong supporter of the genetic mechanism involved in the Darwinian natural selection. By 1917, however, Huxley felt compelled to return to England to enlist in the war. He left Muller in charge and Muller hired Edgar Altenburg to share the teaching while they carried out research on mutation rates and the mutation process. Muller also continued with Altenburg a long-term project started in 1912 that was not published until 1920: a study of what Muller called the gene-character problem.

On theoretical grounds Muller had argued that Darwinian variation has a genetic basis and there must be subtle modifier genes involved in the different expressions of a genetic trait. Morgan had found two such mutants, one called Truncate and the other called Beaded wings. In both cases the shape of the wing varied and neither stock could be made homozygous. They kept throwing off normal winged flies. Morgan had turned these over to Muller to play with and as the years went by, Muller accumulated the evidence that these were complex hereditary systems. In Beaded wings the dominant chief gene (Beaded) was made perpetually heterozygous with rare or no normal-winged flies. The stability of the heterozygote arose from Muller’s determination that Beaded, while dominant for its visible effect, was recessive for a lethal trait. In the homozygous state it killed the embryo. In one line of Beaded a second recessive lethal arose, but it arose in a homologous chromosome, rendering the unrelated lethal perpetually heterozygous. This new lethal was stabilized by a repressor of crossing over (then called a C factor and later recognized as a chromosomal rearrangement, an inversion) (1918).

The analysis of Truncate wings (jointly done with Altenburg) was even more complex, with isolated and mapped modifier genes, which Muller called intensifiers and dimin10 ishers, affecting the expression of the dominant chief gene (1920,1). (Like Beaded, Truncate was a recessive lethal and dominant visible mutation.) There were also environmental modifiers, especially temperature: the mutant expression enhanced at higher temperatures and the normal phenotype at the lower temperature. Muller considered these experiments of supreme importance for the emerging neo- Darwinism, which attempted to bring classical genetics into Darwinian natural selection. In nature, he argued, genes evolve through systems of modifier genes that eventually become homozygous and stabilize the new trait. He also used his analysis to discredit DeVries’s mutation theory. Muller argued that the new species DeVries obtained in Oenothera were actually complexes of chromosomal rearrangements that underwent occasional crossing over, chromosome doublings, or losses of chromosomes, leading to the expression of many changes in the plant’s phenotype and rendering it incapable of breeding with the original type. Oenothera, he argued, was an aberrant mechanism of evolution and not in the Darwinian mainstream.

Muller’s views, initially formulated in the debates with students in the biology club at Columbia, were initially based on theoretical considerations, and he clashed in print with Castle over Castle’s interpretation of hooded rats and other variable traits in small mammals (1914). Castle argued that the genes themselves varied through contamination in the heterozygous state, a claim Muller challenged in his own dissertation work and that he could now demolish with the clear evidence of modifier genes and a reductionist explanation of DeVries’s own competitive model of evolution. For Muller the gene was stable until it was itself mutated and that mutation rate, as he and Altenburg demonstrated, was relatively rare.

Muller left Rice to serve as an interim professor at Columbia while Morgan was away on sabbatical leave. Muller hoped to join the faculty there but Wilson felt it would not work out if Morgan came back, which he did for a few years before leaving to head and develop the new Department of Biology at California Institute of Technology. While at Columbia (1919-1921), Muller published several theoretical papers that charted a future course for his research (1920,2; 1921,1,2; 1926). He argued that mutation should be limited to changes in the individual gene and that they should not be lumped together with other hereditary changes such as nondisjunction, polyploidy, and chromosome rearrangements. He also believed genes should be analyzed through their mutations. He considered the gene as having a unique property in replicating its variations and that something basic was present in the gene, which made it unique to all life forms. He also recognized a similarity between genes and viruses, comparing viruses with “naked genes.” He believed the gene would someday be accessible to chemical and physical analysis.

GENETICS AT THE UNI VERSITY OF TEXAS

Muller returned to Texas but not to Rice. Instead he chose a position at the University of Texas at Austin. He had a powerful influence on his colleagues, especially John Thomas Patterson and Theophilus S. Painter. Patterson was studying armadillo embryology and Painter was working on the cytology of spiders when Muller joined the faculty. After Muller showed the versatility of Drosophila as a tool for genetic analysis, both Patterson and Painter switched their organism of choice and became major contributors to Drosophila genetics. This was both a benefit and a difficulty. It increased the stimulation of discussions and approaches to work in classical genetics, but it also led to a renewed rivalry, with Muller feeling that much of his time and ideas were entering the work of his colleagues and his own work was suffering from neglect. He tried to solve this by working at night, not a very good idea for a married man, and soon his marriage was foundering and, of course, he had alienated Patterson and Painter.

Muller alienated his colleagues as well as the university in other ways. He became an ex officio adviser of the National Student League, named by the FBI as a communist student organization, and he became an underground editor of The Spark, a newspaper promoting socialist goals, including civil rights for African Americans, equality of opportunity for education and careers for women, unemployment insurance for the unemployed, social security for the retired, and other progressive legislation championed especially by the Communist Party, for which Muller had strong sympathies although he never joined.

It is remarkable that as Muller’s personal life became more complicated with marriage to Jessie (who was fired from the faculty in Mathematics when she became pregnant) and conflicts emerged with his colleagues, he became more intensely involved in his major discovery. When he came to Texas he was hoping eventually to induce mutations. He had tried, unsuccessfully, a number of chemical approaches based on the finding that temperature increased the mutation rate in a way consistent with the Q10 of chemical reactions. In 1926 he reexamined the use of radiation. Morgan, Blakeslee, and Payne had tried radiation without success about 1910. Muller realized that a subjective search for mutations was not reliable. (Although Lewis J. Stadler used that method successfully with maize, his papers appearing several months after Muller’s Science paper appeared [1927,1]). Instead Muller designed tools to isolate the most commonly occurring mutations, recessive lethals (first discovered by Morgan).

One of Muller’s great contributions to genetics was stock design. He used complex rearrangements and combinations of recessive and dominant visible markers to identify the passage of chromosomes from parent to offspring. One such stock, called ClB, consisted of a recessive lethal, a crossover suppressor, and the dominant visible mutation called Bar eyes, all on the X chromosome. By irradiating normal or wild-type male flies and having the X chromosomes of their sperm individually rendered heterozygous with the ClB chromosome, Muller could test for the presence of an induced recessive lethal mutation by looking for the absence of that category among the progeny (the grandsons of the irradiated male). This gave Muller a quantitative measure of induced mutations, and he was surprised and elated by an abundance of induced mutations that were 150 times more plentiful than spontaneously arising mutations. He not only obtained the lethal mutations he expected but also visible mutations that were both new and allelic to spontaneous forms previously picked up over the prior 15 years in laboratories around the world.

Muller published his results (without data) in Science (1927,1) to establish his priority and that same year presented the data in great detail at the International Congress of Genetics in Berlin (1927,2). The publicity for Muller’s report of artificially induced mutations was worldwide and Muller returned to the United States with international stature. The Berlin paper mapped the lethals and visible mutations, eliminated competitive models of genes as bean bags of particles, and revealed that a portion of the first generation of mutations was fractional or mosaic (a condition associated with the DNA double helix model and not successfully interpreted until the 1950s).

PERSONAL SUCCESS AND FAILURE AT TEXAS

Muller’s troubles at Texas intensified, as he and Jessie considered separation and Patterson and Muller were no longer on speaking terms. Muller had also intensified his left-wing behavior by bringing two Soviet students to his laboratory, Solomon Levit and Isador Agol, on Rockefeller scholarships. Levit collaborated with Muller on human genetics, a field that Muller felt needed some basic science to improve its study of human traits. Muller had published a paper (1925) on identical twins raised apart arguing that very little was known of the genes involved in human behavioral traits and how they interacted with the environmental factors. He did believe such analysis, like his earlier work with Beaded and Truncate, was eventually feasible, and identical twins was one way to start.

In 1932 Muller’s personal life began to collapse over his marriage, his discontent with Texas, the investigations of the FBI, veiled references to him in the local newspapers as a communist subversive on campus, and claims Stadler and others were making that X rays did not induce gene mutations (as changes in the individual gene) but instead induced chromosome rearrangements of various kinds and sizes. Muller disappeared from his laboratory and did not show up to class, and after his wife called anxious about his whereabouts, a search posse of faculty and graduate students went looking for him in the woods near the outskirts of Austin. He was found walking, muddied, and wrinkled by an overnight rain, and somewhat confused. He had slept off an overdose of barbiturates in a suicide attempt but returned to his class the next day as if nothing had changed.

The suicide attempt occurred just before he left to make presentations at the Third International Congress of Eugenics at the American Museum of Natural History in New York City and at the Sixth International Congress of Genetics in Ithaca at Cornell University. To the amazement of many who knew him, both were major papers that had significant impact on those who heard them. At the eugenics congress Muller presented “The Dominance of Economics over Eugenics” (1932,1). Although C. B. Davenport had attempted to block presentation of the paper, Muller prevailed in his denunciation of the American eugenics movement.

Muller argued that only in a socialist country, where all children, male and female, white and black, had equal opportunities for education, housing, and other social services, would there be an opportunity for a successful eugenics program. American eugenics, he argued, was based on the false premises that social traits such as pauperism, vagrancy, feeblemindedness, and recidivist crime were largely innate traits. Muller argued that this was unproven and probably false. It was a shock to readers of the New York Times (and newspapers around the world) to hear an appeal for the end to racial discrimination, to class-based claims of inferiority, and to the oppression of women. Muller’s phrasing was Marxist and his sympathies with the Soviet Union as the only country where that potential existed (so he believed) was considered both outrageous and reckless for a professor from Texas.

At the Ithaca congress Muller presented a lengthy paper (1932,2), “Further Studies on the Nature of Gene Mutation,” on what might be considered a capstone of classical genetics. Muller presented a theory of gene function in which he introduced the terms “hypomorph” (less than normal activity), “amorph” (no activity at all of normal function), and “neomorph” (brand-new traits that have no counterpart in their normal allelic source). Alleles like apricot or eosin in the white-eyed series were hypomorphs; white itself was an amorph; and the Bar mutation was a neomorph. Muller demonstrated these functions using deleted X chromosomes carrying extra doses of the gene being studied.

Muller also introduced a second discovery. He used fragments of X chromosomes to identify a special category of modifier genes that he called dosage compensators to explain a phenomenon he called dosage compensation in which most genes on the X produce the same outcome quantitatively and qualitatively for gene action whether present in two doses in the homozygous female or one dose in the hemizygous male. Many of those who heard Muller’s presentations were stunned by his originality, his forcefulness in presenting his views, and the importance of what he presented. Others, aware of the rumors surrounding his mental collapse and suicide attempt, found the presentations so incomprehensible and incoherent that they could not take in the importance of what he presented.

Muller returned to pack up and leave for Berlin. He had been awarded a Guggenheim Fellowship to study at the Institute for Brain Research, part of the Kaiser Wilhelm institutes. There he collaborated with N. V. Timofeef-Ressovsky in studies on target theory and the expression of partial dominance by recessive lethals. He arrived in 1932 but the following year Adolf Hitler was elected chancellor and the institute was vandalized by Nazis, who looked with suspicion on the communist leanings of that unit. Muller left to accept an invitation from N. I. Vavilov to come to the Soviet Union and establish a genetics laboratory in Leningrad. At the time, Vavilov’s position was similar to that of the secretary of agriculture in an American president’s cabinet.

Muller’s five years in the Soviet Union (1932-1936) were transforming. He had the best support for his research as corresponding member of the U.S.S.R. Academy of Sciences. He was also free of teaching duties. He built a laboratory first at Leningrad and then at Moscow, where he recruited several graduate students and research associates. The projects he initiated focused on gene function explored through position effect; gene evolution studied through the Bar case; and gene structure analyzed through what he called the left-right test.

In the first of these he noted that certain genes, like scute 19, could be shifted to another chromosome and still retain the original function. Other genes in the region of the tip of the X chromosome, when juxtaposed against heterochromatin showed classical position effect variegation or loss of function (1935 ). The genes themselves, as his students showed, could be isolated by crossovers and restored to normal function. In the Bar case Muller made use of Painter’s discovery of salivary gland chromosomes (a discovery Muller considered so important that he nominated Painter for election to the National Academy of Sciences despite his personality clashes with him back at Texas (1936,1)).

With Alexandra Prokofyeva as his cytologist in the second project Muller showed that the Bar mutation was actually a duplication and he interpreted this as a primary unequal crossover. Once established, Bar tended to revert to normal or produce a triplication, called ultraBar. Muller called this secondary unequal crossing over. The first event Muller associated with extension of individual or small numbers of genes into chromosomes and genomes in the evolution of life from the first gene, and he modified the cell doctrine with what could be called a gene doctrine, which asserts that all genes arise from preexisting genes. Muller’s insight into gene evolution was amply confirmed by the nests of duplicated genes associated with the human hemoglobin A and hemoglobin B genes, each a consequence of extensions by unequal crossing over.

In the third of these projects Muller used independent inversions with breaks, one near the scute region, and the other toward the centromere heterochromatin. These heterozygous inversions provided opportunities to combine the fragments of the yellow-achaete-scute region near the distal tip of the long arm of X chromosome. Muller’s analysis revealed discrete breakage regions between genes, as if there were some inert or nonfunctional material between individual genes (1940).

Muller had ambitious plans for analysis of the gene through radiation-induced mutations and cytological studies. He also was consultant with Levit for the first medical genetics research laboratory. This was constructed in Moscow and included dozens of identical twins that were studied for their physical traits, susceptibility to tuberculosis and other diseases, behavioral responses to mechanical tasks, and similar studies that attempted to sort genetic from environmental factors. The institute published a journal of human genetics (four issues were produced). Muller looked on this pioneering effort as a prelude for his own eugenic ideals. He went ahead with the publication of a book he had started in 1919, which he called Out of the Night (1936,2). He asked Levit’s advice on how a eugenics program could be launched in the Soviet Union, and Levit, a party member, advised him to go to the top.

Muller had the book translated and presented to Premier Stalin with a lengthy letter advocating his utopian dream of positive eugenics in a classless society. It was the wrong time and the wrong idea. At the same time as Muller was hoping to expand his genetic and eugenic programs, a countermovement was underway in Soviet science. Trofim D. Lysenko in Odessa was offering a different view of heredity based on his plant-breeding experiments. He felt that the heredity of a species was malleable if it was shattered by a provocative environment and retrained in the desired direction. Lysenko based his theory on the work of I. V. Michurin, a Russian Luther Burbank, who like Burbank believed Lamarckian transformations by the environment were assimilated by the plants he studied. Michurin’s work was primarily in fruit trees and based on grafting experiments. Lysenko’s work was primarily based on changes in cereal crops, with claims that cold shocks (vernalization) could transform winter into spring wheat or even wheat into rye or oats. Lysenko and his supporters extended their theories to all of heredity, and looked upon western genetics or Mendelism-Weismannism- Morganism as an imported bourgeois capitalist, pseudoscientific system intended to check progress, support racism, and promote fascism.

A bitter debate emerged, with growing support for the Lysenkoists, who successfully lobbied to prevent the 1937 International Congress of Genetics from being held in Moscow. Muller was drawn into the controversy, complicated by the 1936 purge that Stalin had begun through assassinations, arrests, staged trials, and imprisonments of those he considered untrustworthy. Both Agol and Levit were arrested, charged with being Trotskyites, and executed. Muller debated Lysenko in Moscow in December 1936, accusing him of practicing the equivalent of shamanism instead of science and called him a fraud. Muller was shouted down in the uproar at this mass meeting of 3,000 geneticists and collective farmers, about equally divided in their support for genetics or Lysenkoism. Muller realized there was little hope for continued research in the Soviet Union, and Vavilov advised him to find a safe way out. Muller chose to enlist as a volunteer in the Spanish Civil War and he joined the International Brigade, serving with the Canadian physician Norman Bethune doing physiological research on blood transfusion.

THE EDIN BUR GH YEARS

Muller stayed in Spain through the siege of Madrid and when the cause of the Republican Army seemed on the verge of defeat, he tried to find a place to go. He could not return to the Soviet Union, where he would be subject to intimidation, arrest, or execution. He could not return to Austin because he received notice that he would first have to stand trial in the faculty senate for violating a university policy as an editor of an unauthorized newspaper (university policy required signed editorials and columns on student publications). Muller hoped to find work in Paris with Joliot Curie or in Stockholm with Gunnar Lundberg, but they had no openings. Huxley heard of his difficulties and contacted F. A. E. Crew, the director of the Institute for Animal Genetics at the University of Edinburgh.

Crew arranged for Muller to be a guest investigator and Muller found himself once more with an opportunity to develop a graduate program. He arrived in 1937 and he quickly began research with some new problems to examine. He looked at the relation of radiation dose to mutation frequency and with S. P. Ray-Chaudhuri demonstrated that the same amount of mutation is produced by a given dose whether that dose is administered over a month (a protracted dose) or over 30 minutes (an acute dose) (1939). This led Muller to argue that even diagnostic doses of radiation were of concern for radiation protection and he advised practitioners of the danger possible in his annual report to the granting agency that supported his research. Physicians objected that Muller’s views were injurious to patient confidence and inappropriate because the work was done on fruit flies and not human patients. It was the beginning of a skirmish on radiation safety that would persist for the rest of Muller’s life.

Muller had two additional students whose collaboration proved fruitful. With Guido Pontecorvo, Muller worked out ways to use triploid D. melanogaster females and heavily irradiated D. simulans males to construct diploid surviving embryos that carried all D. melanogaster chromosomes except for a fourth chromosome from D. simulans. The analysis of these flies allowed Muller and Pontecorvo to argue that interspecific hybrids that fail to survive owe their failure to assignable genetic differences rather than to some vague mixing of incompatible cytoplasm. Muller and Pontecorvo also used irradiation and triploid flies to identify the mechanism of dominant lethals. These were aborted embryos produced by radiation exposure of sperm (1942), and Muller and Pontecorvo showed (independently of Barbara McClintock’s work on maize) that dicentric chromosome formation (what McClintock called the breakage-fusion-bridge cycle) was the source of cell death leading to the aborted embryos.

The second student, Charlotte Auerbach, like Pontecorvo was a refugee from fascism. Crew assigned her to Muller. Muller suggested to her that a productive way to study the gene was through mutation and he recommended looking at chemical mutagenesis. Auerbach used pharmacologist Robson’s suggestion to use mustard gas and the first demonstration of a potent chemical mutagen was successfully published (but had to wait until the end of the war because of secrecy laws imposed on agents that were used or could be used for warfare). Also at Edinburgh, Muller met and married his second wife, Thea (Dorothea Kantorowicz). Muller had the frustrating duty of being a leading planner of the aborted Congress of Genetics in Moscow and the transferred congress that met in Edinburgh on the eve of World War II . The outbreak of war put an end to basic research in Great Britain, as a fight for survival dominated all other issues. Muller was advised to move back to the United States.

The best Muller could salvage was an interim position at Amherst College. He did not have the financial support for research, and it was difficult to find assistants willing to work in jobs that were unrelated to the war effort. It was also a time for happiness and rediscovering family life with the birth of his second child, Helen. Muller’s major activities at Amherst were writing review articles. He also worked as a consultant on radiation genetics projects for the then-secret Manhattan Project, but those could not be published. It also meant a return to teaching but Muller’s heart wasn’t in teaching undergraduates. As the war came to an end Muller knew he would not be added to the faculty. He wrote in desperation to friends. McClintock said a letter to her was so alarming in his despair about continuing in academic life that she burned it. Fortunately, Indiana University heard of Muller’s difficulties. Fernandus Payne, who admired Muller’s work, sent Tracy Sonneborn to a meeting to explore Muller’s interest in joining the staff. Muller was delighted, and in 1945 he moved his family to Bloomington.

THE INDIANA YEARS

Muller spent his happiest years in Bloomington. He felt warmly appreciated by his colleagues. He was generously supported by the Rockefeller Foundation and by Indiana University with grants to begin another graduate program. He taught at the graduate level (three courses a year), and he felt vigorous at the age of 55 . In 1946 he was awarded the Nobel Prize, and that had a transforming effect on his position in the university and in national life. It was the third nomination for Muller. The rule of three prevented Muller (as well as Sturtevant and Bridges) from receiving the Nobel with Morgan in 1933 . Lancelot Hogben was asked to write a nomination for Muller in 1939, but war broke out and Muller’s candidacy was deferred. The bombings of Hiroshima and Nagasaki had changed the relation of science to society. An Atomic Age required public debate and Muller’s prize was seen as a message to him and to science to steer society through potential abuse or calamity.

While Muller wore the mantle of elder statesman for science, he was also committed to his graduate program. He studied a variety of projects in radiation genetics using neutrons and other particles, often in collaboration with facilities at Brookhaven National Laboratory. He also looked at new problems in human genetics. He shifted from twin studies as a tool to understanding and reexamined the survival of genes in populations. He made use of his Soviet-period research on the partial dominance of recessive lethals (work done with Kerkis) and extended it to population genetics, using a modification of equations first used by Danforth. Muller presented a new concept that he called genetic load (1950). He believed that spontaneous mutations accumulated in populations and reached an equilibrium with the amount of newly arising mutations matching those eliminated through their partial dominance. In human populations, he argued, the mutational load increases each generation because the pressure of natural selection is relaxed in an unnatural environment.

Muller and his students studied spontaneous mutation rates and used protracted and acute doses under varied physiological conditions (nitrogen- or oxygen-rich atmospheres) that might diminish or enhance chromosome breakage or gene mutation. He refined the tools for genetics and launched numerous stocks to improve the detection of lethal mutations (recessive and autosomal), sterility mutations, and visible mutations.

Throughout those years he was also embroiled in Cold War conflicts. He spoke out against radiation abuses. He was distrusted by those who misinterpreted Muller’s concerns about radiation hazards in medicine, industry, and weapons testing as attacks on national defense and the survival of the West against Stalinist imperialism. He went public on the Lysenko affair after 1948 with an attack on Soviet genetics. At the International Genetics Congress in Stockholm the eastern bloc delegates walked out when Muller started listing the crimes against science he had witnessed in the Soviet Union. Muller was called to testify before the House Un-American Activities Committee. (That testimony is still immune from access). It was an era of fear. He and his wife burned thousands of items they had accumulated in his travels, including correspondence with known communists or communist sympathizers. He hoped to protect his students and colleagues, who like him erred in thinking that the Soviet experiment was democratic and just and free of prejudice.

Muller tried to separate the politics of the Cold War from the very real issues that he felt had to be addressed. What should the maximum exposure doses be for a lifetime of medical diagnosis? How should standards be set for maximum permissible doses into the environment or in the workplace for the nuclear power plants industries were planning? When should scientists support efforts to ban atmospheric and oceanic testing of nuclear weapons? Muller’s views were complex and often misconstrued. He wanted both atomic and hydrogen bombs to be developed. He felt nuclear disarmament was not possible unless it was by mutual agreement in treaties with guaranteed scientific inspection to prevent cheating. He argued that fallout doses (except for the largest of the hydrogen bombs used) were too small to be a public health threat. He argued that diagnostic doses were individually low risk, but when given to hundreds of millions of people, did induce a predictable number of leukemias, solid cancers, and mutations. He argued that the Atomic Bomb Casualty Commission in Hiroshima and Nagasaki would find few mutations in the children of the exposed population because most mutations are recessive and they would not show up for many generations to come. Muller’s views were often rejected by those who feared any dose of radiation however small and by those who dismissed low doses as harmless or even beneficial to the public (because they allegedly created a hybrid vigor in the offspring).

In 1957 Muller revived his eugenic ideals (1959, 1961). He felt the abuses of Nazi eugenics and the American eugenics movement were historical accidents not likely to be repeated in democratic societies. He urged in such societies the adoption of his idea, germinal choice, which should give to the user the decision making on what sperm or eggs to use for producing children. He hoped people would learn to separate sexual activity from the quality of children they desired, just as they had learned to separate sexual activity from reproduction by the adoption of artificial means of birth control. In addition to family planning based on thoughtful desires for children, he recommended a genetic enlightenment that would be most likely to produce healthier, wiser, and more caring offspring. He was criticized in editorials as being ignorant of the Holocaust and the excesses humans are cable of applying against humanity. Many thought he was trying to revive the old-line eugenics he had condemned. Muller realized as his health began to fail that no eugenics was better than bad eugenics, and he refused to endorse a planned sperm bank in California for germinal choice that was based on the values of old-line eugenics.

MULLER ’S LEGACY

Muller led a flawed life. His political involvement in the uses of genetic knowledge made him vulnerable to controversy and negative assessment. It is difficult to speak out on important issues without experiencing rejection or being misconstrued. He told his students that it was their duty to bear witness and to speak out against the abuses of science in their generation. Most scientists have difficulty playing the role of a gadfly. They enjoy doing their science and not worrying about the way their findings will be used. Muller was not alone in taking public stands. Julian Huxley, J. B. S. Haldane, Linus Pauling, Joshua Lederberg, and James Watson are among the many scientists of stature who have spoken out against public policies that seemed injurious to the public. He argued that genetics was the most subversive science because it dealt with issues fundamental to human nature. A geneticist cannot expect to be ignored by those who reject natural selection and evolution. Geneticists are bound to encounter public controversies as their findings are applied to human reproduction. At worst, the government, as in the Soviet Union or in Nazi Germany, may endorse a spurious science to counter the findings developed by geneticists. Muller’s views on eugenics are complex. In the long run he may turn out to be prophetic and genetic-load concerns in distant generations may lead to germinal-choice reproductive options to reduce that load. Muller served humanity well in promoting radiation safety and helping to curb the most egregious abuses of radiation in industry and health.

Muller’s roles in contributing to classical genetics, in founding the field of radiation genetics, and in relating genetics to evolution are solid contributions that will endure. His influence on the careers of many of his colleagues and those who took his courses was profound. He had some successful students, including Bentley Glass, who was elected to the National Academy of Sciences. Both Pontecorvo and Auerbach became fellows of the Royal Society. Many of his students entered academic life and had productive careers. Many of his students in the Soviet Union were not so fortunate, and they spent years isolated from publishing, forced out of genetics, imprisoned, or executed. It is a tribute to Fernandus Payne that he recruited Muller. Payne dismissed the claims of psychosis, communism, and a personality likely to be disruptive to colleagues. Muller had his difficult moments at Indiana but more often than not he brought glory to the university; he respected his colleagues (refusing to teach less than they); and he deeply appreciated the gift of tranquility bestowed on him. Those students who worked with him will appreciate his kindness in encouraging their careers, helping them financially through hard times, and fighting passionately with them on every sentence they wrote in their articles with the conviction that what they did mattered and deserved the tough evaluation of his considerable knowledge.

HONORS AND DISTINCTIONS

In addition to his Ph.D. in 1916 at Columbia University, Muller was the recipient of five honorary degrees: D. Sc., University of Edinburgh (1940); D.Sc., Columbia University (1949); D.Sc., University of Chicago (1959); M.D., Jefferson Medical College (1963); and Ph.D., Swarthmore College (1964). He received numerous prizes and recognitions of his stature in his career: the Cleveland Research Prize, American Association for the Advancement of Science (1967); Nobel Prize in Physiology or Medicine (1946); president, VIII International Congress of Genetics, Stockholm (1948); Kimber Award in Genetics, National Academy of Sciences (1955 ); Virchow Medal, Virchow Society of New York (1956); vice president, International Congress of Radiation Research, Burlington, Vermont (1958); Darwin Medal, Linnaean Society, London (1958); Darwin Medal, Deutsche Akademie Naturforscher Leopoldina (1959); Alexander Hamilton Award, Columbia University (1960); Humanist of the Year, American Humanist Association (1963); and City of Hope Medical Center Research Citation (1964).

Muller was a member of numerous learned societies in the United States, including the National Academy of Sciences (elected in 1931); fellow, American Association for the Advancement of Science; American Society of Naturalists (president, 1943); American Philosophical Society; American Academy of Arts and Sciences; American Society of Zoologists; Genetics Society of America (president, 1947); American Genetic Association (vice president, 1959); Society for the Study of Evolution (president, 1957); American Society of Human Genetics (president, 1949); Society for Experimental Biology and Medicine; American Humanist Association (president, 1956-1959); honorary member, American Institute of Biological Science.

Muller was also elected to the following foreign learned societies: the Royal Society, London; U.S.S.R. Academy of Sciences, corresponding member (1933 , resigned 1948); Royal Danish Academy; Royal Society of Edinburgh; Royal Swedish Academy; Accademia Nazionale dei Lincei; National Institutes of Sciences of India; Akademie der Wissenschaften und Literatur, Mainz; Genetics Society, Japan; Genetical Society; Mendelian Society of Lund; Deutsche Akademie Naturforscher Leopoldina; Japan Academy; Zoological Society, Calcutta; Societa Italiana di Genetica Agraria; Rationalist Press Association; World Academy of Arts and Science (vice president, 1964).

NOTES

1. F or a history of classical genetics see A. H. Sturtevant, A History of

Genetics, New York: Harper and Row, 1965; E. A. Carlson, Mendel’s

Legacy: The Origin of Classical Genetics, New York: Cold Spring Harbor

Laboratory Press, 2004; and J. Schwartz, In Pursuit of the Gene:

From Darwin to DNA, Cambridge: Harvard University Press, 2008.

2. F or a biography of H. J. Muller’s life see E. A. Carlson, Genes,

Radiation, and Society: The Life and Work of H. J. Muller. Ithaca:

Cornell University Press, 1981.

3. M uller’s papers are mostly stored in the Lilly Library at Indiana

University in Bloomington. A smaller collection of Muller correspondence

and papers is stored at the archives of the Cold Spring

Harbor Laboratory Library, New York.

4. F or a biography of Morgan’s life see G. Allen, Thomas Hunt Morgan:

The Man and His Science. Princeton, N.J.: Princeton University

Press, 1978.

5. H . DeVries. Die Mutationstheorie (2 volumes). Leipzig: Viet and

Company, 1901-1903.

6. F or two different views of this group dynamics see E. A. Carlson,

The Gene: A Critical History, Philadelphia: Saunders, 1966; and J.

Schultz, Innovators and Controversies, Science 157(1967): 296-301.

For Sturtevant’s view see his chapter “The Fly Room” in A. H. Sturtevant,

A History of Genetics, New York: Harper and Row, 1965.

7. T he mechanism of crossing over. Parts 1-4. Am. Nat. 50:193-221,

284-305, 35 0-366, 421-434.

SELECTED BIBLIO GRAPHY

For a complete listing of Muller’s 372 publications and a selection of complete

and partial works, see H. J. Muller, Studies in Genetics, The Selected Papers

of H. J. Muller, Bloomington: Indiana University Press, 1962.

1914

The bearing of the selection experiments of Castle and Phillips on

the variability of the gene. Am. Nat. 48:567-576.

1916

The mechanism of crossing over. Parts 1-4. Am. Nat. 50:193-221, 284-

305, 35 0-366, 421-434.

1918

Genetic variability, twin hybrids and constant hybrids in case of balanced

lethal factor. Genetics 3:422-499.

1920

[1] With E. Altenburg. The genetic basis of truncate wing: An inconstant

and modifiable character in Drosophila. Genetics 5:1-59.

[2] Further changes in the white eye series of Drosophila and their

bearing on the manner of occurrence of mutations. J. Exp. Zool.

31:443-473.

1921

[1] Mutation. The Third International Congress of Eugenics. In

Eugenics, Genetics, and the Family 1923 pp. 495-502.

[2] Variation due to change in the individual gene. Am. Nat. 56:32-50.

1925

Mental traits and heredity as studied in a case of identical twins

reared apart. J. Hered. 16:433 -448.

1926

The gene as the basis of life. Proceedings of the International Congress

of Plant Sciences 1:897-921.

1927

[1] Artificial transmutation of the gene. Science 66:84-87.

[2] The Problem of Genic Modification. Verhandlung die V Kongres fur

Vererbungslehre: Suppll. Bd. I des Zeitschrift fur inductive Abstammungs

und Vererbungslehre, pp. 234-260.

1932

[1] The dominance of economics over eugenics. Sci. Mon. 37:40-47.

[2] Further studies on the nature and causes of gene mutations. Proceedings

of the 6th International Congress of Genetics, Ithaca 1:213-255 .

1935

The origination of chromatin deficiencies as minute deletions subject

to insertion elsewhere. Genetica 17:237-252.

1936

[1] Bar duplication. Science 83:528-53 0.

[2] Out of the Night: A Biologist’s View of the Future. New York: Vanguard.

1939

With Ray-Chaudhuri. The validity of the Bunsen-Roscoe law in the

production of mutations by radiation of extremely low intensity.

Proceedings of the Seventh International Congress of Genetics.

J. Genet. Suppl.:246.

1940

With D. Raffel. Position effect and gene divisibility considered in connection

with strikingly similar scute mutations. Genetics 25:541-583.

1942

With G. Pontecorvo. The surprisingly high frequency of spontaneous

and induced breakage and its expression through germinal

lethals. Genetics 27:157-158.

1950

Our load of mutations. Am. J. Hum. Genet. 2:111-176.

1956

On the relation between chromosome changes and gene mutations.

Brookhaven Sym. Biol. 8:126-147.

1959

The guidance of human evolution. Perspect. Biol. Med. 3:1-43.

1961

Human evolution by voluntary choice of germplasm. Science 134:643-

649.