Tag Archives: Franklin Roosevelt

The Remarkable Journey of Medical Genetics Pioneer, Arno Motulsky

Dr. Arno G. Motulsky, recognized for having pioneered the field of “pharmacogenetics”—the genetic basis for differences in the way individuals respond to drugs—passed away on January 17, 2018 in Seattle, at the age of 94. Since Motulsky was not a virologist, his passing is noted here not for his scientific contributions but, rather, for his improbable achievements against very long odds and, especially, for the remarkable journey from which this story began—a journey which brings to mind a current issue before the public. It began as follows.

Dr. Arno Motulsky in his Seattle apartment in 2014. Credit Clare McLean/University of Washington

1938 saw a dramatic increase in violent anti-Jewish acts by the Nazi regime in Germany. In May 1939, 916 passengers, including 910 German Jews seeking to escape from the Nazis, were aboard the SS St. Louis; a liner of the Hamburg-America Line, which was bound for Cuba. Hundreds of thousands of other German Jews would not be able to get out.

Each of the passengers on the St. Louis had a visa from the Cuban government that was issued before the ship departed from Germany. However, unbeknownst to the passengers—but known to the shipping line—the President of Cuba, Federico Laredo, had invalidated the visas the day before the ship sailed. Thus, when the St. Louis arrived in Havana Harbor, the Cuban government would not allow the passengers to disembark. Instead, the Cuban authorities attempted to exploit the refugees’ situation in order to extort bribes from them. These bribes began at $500 a person and rose to $1,000,000! All the while the ship sat within yards of the shore.

Cuba had been practicing this sort of extortion before the incident involving the St. Louis. Manuel Benitez Gonzalez, the Director-General of the Cuban immigration office, routinely sold landing documents for $150 or more and, according to U.S. estimates, had amassed a personal fortune of $500,000 to $1,000,000. Eventually, Benitez’s corruption would lead to his resignation. However, corruption was not the only factor working against the refugees. Many Cubans saw them as competitors for scarce jobs. Moreover, Cuban sympathy for the refugees’ plight was also tempered by xenophobia and antisemitism. [News reports of the impending sailing of the St. Louis triggered anti-Semitic demonstrations in Cuba even before the ship left Germany.]

As the St. Louis sat at anchor in Havana Harbor, the American public raptly followed daily media accounts of the refugees’ plight. And despite urgent appeals from American sympathizers (non-Jewish as well as Jewish), the U.S. State Department and the White House decided not to intervene on behalf of the refugees. President Franklin Roosevelt did not answer telegrams from the refugees on the ship. The State Department did respond, wiring back to the passengers that they must “await their turns on the waiting list and qualify for and obtain immigration visas before they may be admissible into the United States.”

Roosevelt had the power to issue an executive order that would have allowed the St. Louis’ refugees to enter the United States. He had, in fact, permitted 15,000 German and Austrian Jews, who were in the United States on visitor’s permits, to stay, saying: “It would be a cruel and inhuman thing to compel them to leave here” for Germany.   

Why then did Roosevelt and the State Department abandon the refugees on the St. Louis? American popular opinion, which was opposed to accepting more new arrivals, was a key factor. Indeed, about 80% of Americans were against easing immigration restrictions. A key reason was that the depression left many Americans out-of-work. Consequently, many were fearful of foreigners becoming competitors for scarce American jobs. Antisemitism, isolationism (in the years after World War I, many Americans were staunchly against the U.S. being drawn into European affairs), and xenophobia were no doubt factors as well.

The National Origins Act of 1924 permitted only a fraction of would-be emigres from Germany to enter the United States. The quotas that were set on countries were intended to favor “desirable” immigrants from Northern and Western Europe, while limiting less “racially desirable” immigrants, such as Southern Europeans, Eastern European Jews, and people born in Asia and Africa. Yet anti-German sentiment may have led American immigration officials to allow only one fourth of the German quota to be filled. Also note that three months before the St. Louis sailed to Cuba, the U.S. Congress let die a bill that would have admitted 20,000 Jewish children from Germany above the existing quota.

Critics contend that despite the political pressure on Roosevelt, he might have done more to raise public sympathy for the plight of the refugees. Moreover, once the United States was in the war, and the U.S. government became aware of Hitler’s intent to annihilate all Jews in Germany and in German controlled areas, the government tried to withhold that information from the public. And as the facts about the murder of the European Jews—concentration and extermination camps—leaked out, and despite pleas from Jewish leaders in Warsaw and elsewhere in Europe, Roosevelt would not act to impede the extermination program (e.g., by ordering the bombing of Auschwitz and the rail lines to it). Instead, Roosevelt maintained that winning the war was the only way to save the refugees, and winning the war was the absolute priority.

Barriers to Jewish immigrants were also in place across Latin America. [Note that the British refused to consider Palestine as a solution to the Jewish refugee problem.] Thus, Columbia, Chile, Paraguay, and Argentina rejected appeals from the Joint Distribution Committee (JDC, the major Jewish agency for relief overseas) for haven. And, when the JDC could not meet a deadline set by Cuba for the payment of ransoms, the St. Louis was forced to leave Havana Harbor. Remarkably, the German captain of the St. Louis, Gustav Schröder (who had been unaware of the Cuban government’s duplicity before the St. Louis sailed), appealed to the United States for haven. But his personal pleas were unheeded. What’s more, as the St. Louis sailed past Miami and the Florida coast, U.S. Coast Guard ships made certain that none of its passengers might escape to freedom. So, the St. Louis turned back to Europe, with sixteen-year-old Arno Motulsky among its passengers.

The German liner St. Louis being denied entrance to Havana, Cuba, in June 1939. Dr. Motulsky was aboard the ship with his mother and siblings when it was sent back to Europe. Credit Associated Press

But the St. Louis did not return to Germany. The JDC had arranged with Great Britain, France, Belgium, and the Netherlands for each of these countries to take one fourth of the refugees. Still, only the 288 refugees admitted to Great Britain would be safe, since the Nazis would overrun the other countries within months. Only a few would survive the Holocaust.

Motulsky was among the refugees who disembarked from the St. Louis in Belgium. He then spent a year in internment camps in Vichy France, where many other detainees died from starvation or typhoid. He reached the United States in June of 1941; ten days before his 18th birthday. His account of his flight to freedom, in his own words, is as follows:

“…By early 1939, my parents realized we would need to leave Germany. My dad had a brother in Chicago, so we hoped to go there, but in order to get out of Germany and into America, one needed a visa, which required a quota number that took a while to come up. So my father went to Cuba, where immigration permits were available, with the idea that we would join him later in Chicago. But over the next few months, as the situation deteriorated in Germany, my parents decided it would be best for my mother and brother and sister and me to join my father in Cuba and go to Chicago later.

We left Hamburg in May 1939, on a ship named the St. Louis. There were almost a thousand Jewish refugees on board. The voyage from Hamburg to Havana was normal. But before we could disembark in Havana, the Cubans declared our permits void and refused to let us land. The ship was German, but the captain, Gustav Schröder, was very sympathetic to us and sailed all over the Caribbean in search of a friendly port. Of course, we asked to land in America, but were denied. When we sailed close to Miami, US Coast Guard cutters and planes shooed us off.

So ultimately the ship had to return to Europe. I was 16, so still a little naive, but many of the passengers had been interned previously and knew what awaited us on our return. Some attempted suicide by jumping overboard. Telegrams were sent all over the world asking that we be allowed to disembark anyplace other than Germany. Miraculously, a few days before we would have arrived back in Germany, four other countries—England, France, Holland, and Belgium—each agreed to take one-fourth of the passengers. By luck of the draw, my mother and brother and sister and I were assigned to Belgium. So in June 1939, I started high school in Brussels.

Eleven months later, on May 1, 1940, my dad got his visa to move from Cuba to the US and went to Chicago. On the same day, our US visas came through as well. But by May 10, we hadn’t yet gotten out of Belgium, and the Germans invaded. Leaving was impossible. Since I was now 17, I was arrested by the Belgians as an enemy alien—ironically, as a German—and sent to an internment camp in France. As the Germans invaded France, we prisoners were moved further and further south, until we ultimately were interned in a camp in the Pyrenees, in Vichy-controlled France. The Vichy French, who collaborated with the Germans, sent the ethnic Germans back to Germany and kept the Jews interned. The internment camps had no food or sanitation. Many prisoners died, most from typhoid or starvation.

About ten months later, those of us with US visas were allowed to go to another camp, near Marseilles, where there was an American consulate. My visa had expired, but I pleaded for its renewal. It was ultimately granted, and in June 1941, ten days before my 18th birthday, I left France legally, crossed Spain into Portugal, and sailed from Lisbon to America. Ten days later and I wouldn’t have made it, because Franco did not allow males 18 or older to pass through Spain. A few months later, the Vichy French turned over all their internment camps to the Gestapo.

In August 1941, I arrived in America and joined my father in Chicago. Two years later, we learned that my mother and brother and sister had also survived. With the help of Belgian friends, they had crossed illegally into neutral Switzerland, where they were allowed to live unharmed until the end of the war. My family reunited in Chicago in early 1946.”

In Chicago, in 1942, 18-year-old Motulsky was examined on material he learned in the informal classes taught in the internment camps in France, and was then granted a high school certificate. Motulsky then enrolled in evening classes at Central YMCA College (which later became Roosevelt University) in Chicago, while working at a job during the day. He met his future wife, Gretel, in his English class at the college. He notes that Gretel was the better English student because she survived the war in England and America after leaving Germany in 1938. They were married in 1945.

Remarkably, in 1943, Motulsky was accepted by the medical school of the University of Illinois in Chicago. He also became a U.S. citizen, joined the Army, and was assigned to a specialized program at Yale for rapid training of young physicians. During his year at Yale, Motulsky completed the premedical courses that he still was lacking. More importantly, he also “took genetics with Donald Poulson and was hooked forever.”

In 1944 Motulsky served briefly as an orderly at an Army hospital near Boston, before returning to medical school at the University of Illinois, as a private first class. He was released from the Army in 1946, finished medical school in 1947, and began his residency in internal medicine at Chicago’s Michael Reese Hospital; supported by a fellowship in hematology. Under his fellowship mentor, biochemist and hematologist, Karl Singer, Motulsky began his foray into research, investigating hereditary hemoglobin disorders.

In 1951, the Korean War broke out and Motulsky was again in the Army; this time at a new hematology research unit at Walter Reed Hospital in Washington, DC. “At Walter Reed, we studied hemolytic anemias that might explain problems encountered by our soldiers… Singer had encouraged me to think about biochemical mechanisms, and here were genetic diseases, for which mechanisms could be studied to the direct benefit of patients.”

In 1953 Motulsky joined the faculty at to the University of Washington in Seattle, where he taught internal medicine and specialized courses in hematology. That same year, Watson and Crick determined the structure of DNA, rousing Motulsky to slip what he called “bootleg medical genetics” into his hematology lectures. In 1957 he founded the Division of Medical Genetics at UW; one of the first such divisions in the United States. [The Division of Medical Genetics at Johns Hopkins opened the same year.] But, before that, to broaden the scope of his knowledge, Motulsky spent eight months in the human genetics unit of Lionel Penrose at University College London. “Penrose’s department was the best in human genetics in Europe, including, in addition to Penrose himself, J.B.S. Haldane… People were very critical and helped me recognize excellent work.” Concurrently, Motulsky became interested in what is now called pharmacogenetics—”that is, differential reaction to drugs among people based on their genotypes.”

This blog post was meant to recount Motulsky’s remarkable personal journey during the early years of his life and career. Nonetheless, Motulsky’s considerable scientific contributions also need to be noted, if only briefly—in particular, his analyses of the genetic predisposition to a broad assortment of conditions, including heart disease, blood disorders, colorblindness, infections and immunity, hypertension, and alcoholism (for a concise account of his achievements, see his short autobiography referenced below). It is a good story. For now, note the comments of Francis Collins—a geneticist, and the director of the National Institutes of Health—which Collins made after Motulsky’s passing. “The relationship between heredity and the response to drug therapy — nobody was thinking about that until he started, 60 years ago. He anticipated it decades before science made it possible to get the answers that he dreamed of.” Moreover, “There were very few medical centers that considered genetics as being all that relevant to human medicine. It was more a study of fruit flies and mice, not humans.” Also note the comment of Claire King, Motulsky’s colleague at the University of Washington (and reporter of his autobiography), who discovered the role of certain genetic mutations in breast cancer. Because of Motulsky’s work in medical genetics, “the field is now integrated into every other field of medical practice and has become the soul of precision medicine.”

Motulsky ends his autobiography as follows: “Throughout my career, I have very much enjoyed the practice of medicine. Until I was 70 years old, I was an attending physician in general internal medicine. The human contact, to be able to help people, is enormously rewarding. And I always learned something. I was stimulated to ask new questions by seeing things that I hadn’t previously thought about. And with respect to medical genetics in particular, I know of no other field in science and medicine that is as fascinating. Medical genetics is closer to science than most specialties in medicine. It reveals exciting new biological phenomena. It has social implications and historical implications and ethical implications. I cannot imagine having done anything more exciting for the past 70 years.”


Dallek, R. Franklin Roosevelt and American Foreign Policy, 1932-1945. Oxford University Press. 1979.

Berenbaum, M. The World Must Know: The History of the Holocaust as told in the United States National Holocaust Memorial Museum. Little, Brown and Company, 1993.

Gradyjan, D. Arno Motulsky, a Founder of Medical Genetics, Dies at 94. Obituary, N.Y. Times, January 29, 2018

Motulsky, A. G. (as told to Mary-Claire King). 2016. The Great Adventure of an American Human Geneticist. Annual Review of Genomics and Human Genetics 17:1-15. https://doi.org/10.1146/annurev-genom-083115-022528


Several posts on the blog feature individuals whose lives were impacted in major ways by World War II and the Holocaust. In particular:

The Life and Legacy of George Klein: Cancer Virus Pioneer and Witness to the Holocaust, May 4, 2017.

Gunter Blobel: Nobel Laureate Best known For the Signal Hypothesis of Protein Targeting, February 22, 2018

The Converging Lives of Jacques Monod, Francois Jacob, Andre Lwoff, and Albert Camus in Wartime France, March 27, 2017



Dr. Arno Motulsky in his Seattle apartment in 2014. Credit Clare McLean/University of Washington

Julius Youngner: Slighted Polio Vaccine Pioneer

This is a tale of the hurt that a junior investigator might feel when a senior investigator takes the lion’s share of the credit for the junior investigator’s crucial breakthroughs. Jonas Salk, who conceived and oversaw the development of the first widely used polio vaccine, is the senior investigator in this anecdote. Julius Youngner, the last surviving member of the original vaccine research team that Salk assembled in the early 1950s at the University of Pittsburgh, is the slighted assistant. Youngner later had his own distinguished career. He passed away in April of this year. Here is their story.

After earning his Ph.D. in microbiology, Youngner was drafted into the World War II U.S. Army, which assigned him to the Manhattan Project, to test the toxicity of uranium salts. Youngner first learned the purpose of the Manhattan Project when the first atomic bomb was dropped on Japan.

After the war, Youngner worked as a commissioned officer for the U.S. Public Health Service. This was a significant stop in his career, since it was there that he first became interested in viruses and cell culture. But, since there was no opportunity for him to pursue that interest in Bethesda, he began to look elsewhere. Thus, it happened in 1949 that Salk recruited Youngner to join his vaccine research team in Pittsburgh, after a mutual acquaintance told Salk that Youngner was eager to work on viruses and cell culture.

Jonas Salk and Julius Youngner at the University of Pittsburgh, early 1950s

Salk hoped that Youngner might find a way to generate enough cells from monkey kidney tissue to support mass-production of the vaccine. Youngner, on his own, then developed the use of the proteolytic enzyme, trypsin, to disperse tissue fragments into individual cells, thereby generating many more cells from a given amount of tissue. Indeed, Youngner could generate enough cells to support manufacture of the vaccine. This was his first key contribution to the vaccine project. “Trypsinization” remains a mainstay of modern cell culture.

Youngner’s next major contribution to the vaccine enterprise was his development of a rapid analytical test that had two crucial applications. First, recalling that the Salk vaccine contains an inactivated virus, Youngner’s so-called “color test” made it possible to quickly screen batches of the vaccine for any live virus that might have survived the inactivation process.  Second, Youngner’s test made it possible to quickly test the vaccine’s ability to induce anti-poliovirus antibodies (1). [Youngner based his color test on an earlier observation by John Enders, Tom Weller, and Fred Robbins, that metabolic activity (as indicated by a drop in pH) was less in cultures inoculated with live virus than in control cultures (2, 3). In Youngner’s test, a color change of phenol red, resulting from a shift in pH, served as an indicator of virus activity, or of antibody activity.]

Some sources credit Youngner with having devised the process for inactivating the virus. But, that is correct in a very limited sense only. Salk selected incubation in formalin as the means to disable the virus. In truth, Salk learned of that approach a decade earlier while doing postgraduate studies under Thomas Francis at the University of Michigan. Francis was then using formaldehyde to produce his killed influenza vaccine (2).

What’s more, Salk’s choice of formalin to generate his polio vaccine was bold. Earlier, in the 1930s, Canadian scientist Maurice Brodie tested a formalin-killed polio vaccine in twelve children, with disastrous results. Several of the children developed paralytic poliomyelitis (4).

Clearly, too little exposure to formalin could leave enough live virus to cause paralytic poliomyelitis or death. On the other hand, too much exposure could so badly damage the virus’ proteins that they might no longer induce an immune response against the live virus. Brodie did not have analytical procedures to ensure that he had inactivated his vaccine to safe levels. In contrast, it was clear to Salk that getting the correct balance would be vital to his vaccine project, and Youngner’s color test was the means for doing so. Youngner used his test to determine that six days of incubation in a 1:4,000 formalin solution would result in one live virus particle in 100 million doses of the vaccine (5).

Since Youngner’s inactivation curve was based on only a few data points, and since it was likely that the slope of the curve might flatten out after a time, Salk added a margin of safety of six extra days. Thus produced, the vaccine induced antibody production in monkeys, while showing no signs of causing paralysis or other problems.

By 1954, 800,000 children had been successfully immunized against polio in the first clinical trial of the vaccine. In April 1955, the outcome of the trial would be announced to a very grateful public.

By 1957, Salk’s vaccine team at Pittsburgh was no longer needed, and was dispersing. Salk was making plans to leave Pittsburgh for California, where he would found the prestigious Salk Institute. Youngner, now 34 years-old, remained at Pittsburgh, where he would begin his own distinguished career.

Although Youngner was now independent of Salk, he remained bitter over his former boss’s failure to acknowledge the underlings who had labored so diligently behind the scenes to bring the vaccine to fruition. “The first rule we learned was to call him ‘Dr Salk,’ never Jonas. He would speak to us through a wall of notes and memos…Here was a guy who could always find an hour to brief some reporter at the local Chinese restaurant, but could never find the time to sit down with his own people (6).”

Youngner was particularly appalled by events involving the paper he wrote describing his color test. “After I had what I considered to be a good draft…I gave my copy to Jonas for his comments. It should be noted this was 1954, the pre-Xerox, pre-word-processing era. I had made a working transcript of the paper for my own use and it was this copy that I handed to him. Also, it should be noted that the title page had the authors listed as ‘J.S. Youngner and E.N. Ward (6).’” Elsie Ward, who served as Youngner’s technician, was a zoologist who specialized in growing viruses.

Salk intended to read Youngner’s manuscript while away on a trip.  When Salk returned a week later, he claimed that he had lost the manuscript, but that he had jotted down some notes from which he was able to produce a draft of his own. Youngner was rather incredulous that a person as meticulous and disciplined as Salk could lose such an important manuscript. Youngner’s skepticism was further roused by the fact that Salk’s version contained all the data in Youngner’s original manuscript. Salk explained that incongruity, alleging that he found Youngner’s tables, but not the text.

In any case, Youngner was especially upset by a specific change Salk made to the title page of the manuscript: “The authors were now ‘Jonas E. Salk, J.S. Youngner, and Elsie N. Ward.’ When I (Youngner) questioned the change, Jonas said that since he had to reconstruct the whole paper it was only fair that his name go first…It was obvious to me then, and is more so now, that he considered the advance in this paper a major one and he wanted his name associated with it, even though at the time he had done nothing in the lab (no kidding!) or of an advisory nature to initiate or carry out the work (6).”

Youngner could grudgingly accept that project leaders often used their senior position to appear as co-authors, or even principal authors, on papers emanating from their labs, even if their contributions were minimal. What troubled Youngner in this instance was not that Salk pulled rank, but rather his seeming duplicity.

In yet another instance—the 1955 public announcement of the successful outcome of the clinical trial—Youngner again sensed “a pattern of deception on Salk’s part to take undue credit for the discoveries of others (6).” Salk advocated for the announcement to happen at the University of Pittsburgh. However, the National Foundation for Infantile Paralysis (better known as the “March of Dimes”), which funded the vaccine project, chose the University of Michigan in Ann Arbor as the site for the announcement. That was where Michigan professor Thomas Francis supervised the evaluation of the field trial. [Note that the NIH was not able to fund research back then the way it can today. Thus, the polio vaccine project was supported nearly entirely by private donations to the National Foundation.]

Thomas Francis spoke first. Then, when Salk spoke, he acknowledged the more prominent players in the vaccine project, including Thomas Francis, Harry Weaver (director of research at the National Foundation), Tom Rivers (chairman of the advisory committees on research and vaccines for the  National Foundation), and Basil O’Connor (law partner of Franklin Roosevelt, recruited by Roosevelt in 1928 to raise funds for polio patients at Roosevelt’s Warm Springs Foundation, and a co-founder with Roosevelt of the National Foundation in 1938; (2)). Salk then acknowledged various deans and trustees at the University of Pittsburgh. Yet, he made no mention whatsoever of his dedicated coworkers in his laboratory. They had been expecting at least some recognition from their boss.

Some of Salk’s defenders argued that Salk had acted in the best scientific tradition by prefacing his printed remarks with the phrase, “From the Staff of the Virus Laboratory by Jonas E. Salk, M.D.” But, this was small consolation to Youngner and others of Salk’s coworkers, who expected to be individually acknowledged for their exhausting work on behalf of the life-saving vaccine. Indeed, they felt betrayed.

At any rate, the 1955 announcement of the success of the polio vaccine field trials was joyously received by the public. And while Youngner remained embittered over Salk’s slighting of his coworkers, he nonetheless understood that from the point of view of the National Foundation, “it was much easier to continue raising money when you have a hero, and they had an enormous public relations department that took up Jonas’ name as the hero, which he deserved…But in the meantime, Jonas was, how shall I say, not very generous to his colleagues and he made sure that nobody else was ever mentioned (6).”

The following excerpt is from Polio: An American Story (6). “In September 1963, Salk returned to Pittsburgh to attend the unveiling of his portrait in the auditorium of the University’s medical complex, a stone’s throw from the hospital where he had done his historic polio research. Before the ceremony, Salk told Dean George Bernier that he wished to speak privately with his former assistant, Julius Youngner, now a distinguished professor at the school of medicine. The two men hadn’t talked or crossed paths since Salk’s move to California in 1961. Salk saw the meeting as a courtesy to the only remaining member of his laboratory staff; Youngner had a different agenda. Speaking softly, he recalled, he slowly released the ‘hurt’ he had bottled up for more than thirty years. ‘Do you still have the speech you gave in Ann Arbor in1955? Have you ever reread it?’ Youngner began. ‘We were in the audience, your closest colleagues and devoted associates, who worked hard and faithfully for the same goal that you desired…Do you remember who you mentioned and who you left out? Do you realize how devastated we were at that moment and ever afterward when you persisted in making your coworkers invisible? Do you know what I’m saying,’ I asked. He answered that he did…Jonas was clearly shaken by these memories and offered little response.’…The two men engaged in some uncomfortable small talk before Dean Bernier returned to escort them to the ceremony. Speaking later to a reporter, Youngner admitted, ‘I got a lot of things off my chest. I’m beyond the point where I pull my punches with him. I think it was the first time he ever heard it so graphically.’ Asked if he had any regrets about working for Salk, Youngner replied: ‘Absolutely not. You can’t imagine what a thrill that gave me. My only regret is that he disappointed me.”’


Jonas Salk is deservedly celebrated for developing the killed polio vaccine. That vaccine, together with Albert Sabin’s live attenuated vaccine, which followed soon afterwards, has nearly eradicated polio worldwide. Importantly, Sabin and other polio researchers believed that only a live vaccine could induce a level of immunity sufficient to protect against a challenge with live virulent virus. Nonetheless, Salk persevered in his conviction that a killed vaccine could protect against polio, and he was right.

Salk founded the prestigious Salk Institute in 1963. Yet he never himself made another notable contribution to science.

Youngner may be best known for his work on the Salk vaccine. Yet he had a distinguished career of his own at the University of Pittsburgh after Salk left. Youngner is especially noted for his contributions to interferon research. These include his finding that non-viral agents could trigger interferon induction in animals. And, in collaboration with colleague Samuel Salvin, he identified a second type of interferon, now known as gamma-interferon. Youngner also helped to explain the antiviral-effect of interferon, and he was the first researcher to demonstrate that some viruses express countermeasures against interferon.

Youngner also made important findings in the area of persistent virus infections. Importantly, he demonstrated that defective viral variants, including temperature-sensitive mutants, can play a role in the establishment and maintenance of viral persistence; doing so by impairing (modulating) the replication of the wild-type parental viruses. Based on that principle, Youngner sought to develop dominant-negative mutants of influenza virus as a novel means of anti-influenza therapy. In addition, Youngner and colleague Patricia Dowling developed a novel live attenuated vaccine against equine influenza virus, based on a cold-adapted influenza virus, which can replicate only at the temperatures found in the respiratory tract. That live vaccine was the first to prevent a serious respiratory disease of horses.

Julius Youngner, 2010


  1. Salk, J.E., Youngner, J.S, Ward, E.N. (1954). Use of Color Change of Phenol Red as the Indicator in Titrating Poliomyelitis Virus or Its Antibody in a Tissue–Culture System,” American Journal of Epidemiology. 60: 214–230.
  2. Jonas Salk and Albert Sabin: One of the Great Rivalries of Medical Science, Posed on the blog March 27, 2014.
  3. John Enders: “The Father of Modern Vaccines,” Posted on the blog August 4, 2016.
  4. Vaccine Research using Children, Posted on the blog, July 7, 2016.
  5. Williams, G., Paralysed with Fear: The Story of Polio, Palgrave Macmillan, 2013.
  6. Oshinsky, D.M., Polio: An American Story, Oxford University Press, 2005.

The Converging Lives of Jacques Monod, Francois Jacob, Andre Lwoff, and Albert Camus in Wartime France

In the 1950s and 1960s, two French biologists at the Pasteur Institute, Francois Jacob and Jacques Monod, explained how genes are regulated in bacteria. Their studies of the “lac operon” of E. coli indeed opened up the field of gene regulation, and were a key development in the new science of molecular biology. Their experimental findings also implied the existence of an unstable intermediate between genes and protein synthesis, which eventually led to Jacob’s discovery, in collaboration with Sydney Brenner and Matt Meselson, of messenger RNA (1).

Jacob and Monod shared in the 1965 Nobel Prize for physiology or medicine for their breakthrough studies on gene regulation. Fellow Pasteur Institute scientist, Andre Lwoff, received a share of the award for his pioneering studies on the nature of lysogeny (i.e., how a bacteriophage’s genome can be incorporated into the genome of a host bacteria, and remain latent until being activated by an inducing factor).

In 2013, evolutionary biologist Sean B. Carroll published a book—Brave Genius: A Scientist, a Philosopher, and their Daring Adventures from the French Resistance to the Nobel Prize—that relates how wartime circumstances brought together Jacques Monod and his scientific colleagues Francois Jacob and Andre Lwoff (2). But while much of that story is already known (3), Carroll also tells us of the little known, but remarkable coming together of Monod and philosopher/writer Albert Camus, one of the intellectual giants of the 20th century. Coming from very different intellectual backgrounds, Monod and Camus forged a deep friendship, united in their opposition to tyranny and oppression. Carroll’s book was the inspiration for this post.

When Albert Camus learned that he had won the Nobel Prize for Literature in October 1957, he wrote to a few well-wishers, including an old friend in Paris:

My dear Monod,

      I have put aside for a while the noise of these recent times in order to thank you from the bottom of my heart for your warm letter. The unexpected prize has left me with more doubt than certainty. At least I have friendship to help me face it. I, who feel solidarity with many men, feel friendship with only a few. You are one of these, my dear Monod, with a constancy and sincerity that I must tell you at least once. Our work, our busy lives separate us, but we are reunited again, in one same adventure. That does not prevent us to reunite, from time to time, at least for a drink of friendship! See you soon and fraternally yours.

                                                                                                                                             Albert Camus

Camus appears somewhat downcast in his note to Monod. At 43-years-in-age, he was the second youngest writer ever to receive the Nobel Prize for literature (Rudyard Kipling at 42 was the youngest), and he was worried that the ballyhoo surrounding the award might distract him from his writing. And, he was concerned that the prize might stir up additional contempt from critics of his writing, as well as from his leftist colleagues who opposed his condemnation of Soviet communism.

But, why did philosopher/writer Camus—an intimate of some of the greatest writers and artists of the mid-twentieth century, including Sartre and Picasso—write to scientist Monod, and acknowledge the special importance he placed on their friendship? Likewise, why did he assert: “I have known one true genius, Jacques Monod.” And, what is the same adventure that Camus refers to?

A brief background to our tale is as follows. In March 1939, Hitler took control of Czechoslovakia. Next, on September 1, Germany invaded Poland. On September 2nd, Poland’s allies, Britain and France, issued an ultimatum to Germany: withdraw or face war. On September 3rd, the ultimatum expired, Britain and France declared war on Germany, and the Second World War was underway; sort of. Although Germany went on to conquer Poland in a mere eight days, several months passed without further action. Then, in May 1940, Nazi Germany invaded and overran France in just six weeks. Marshall Pétain surrendered to the Germans, the French Forces were disbanded, the pro-Nazi Vichy government was put in place under former prime minister Pierre Laval and the 84-year-old Pétain, and the Nazi occupation of the defeated French nation began.

A few months before the Nazis invaded France, thirty-year old Jacques Monod was a doctoral student in zoology at the Sorbonne. [A polymath, he also founded a Bach choral group, and was an accomplished cellist, and seriously considered a career in music (4).]  But as war with Germany loomed, Monod enlisted in the army—in the communication engineers—where he thought he might use his scientific talents if war were to break out. Consequently, Monod was serving on the front lines when the Germans invaded. France suffered the most colossal military disaster in its history, and Monod returned to his studies in Paris.

Life in France grew progressively harsher under the Nazis; beginning with subjugation, and followed by deportations, enslavement, and mass murder. Early on, Monod joined one of the first units of the French Resistance; a group of ethnologists and anthropologists at the Musée de l’Homme (Museum of Man).

One of Monod’s duties for the Musée de l’Homme group was to distribute its newspaper, at night. This seemingly simple task was extremely dangerous since capture could mean deportation to a concentration camp or execution. Monod, in fact, had several close escapes. On one occasion, the Gestapo raided his laboratory at the Sorbonne. Fortunately, since they were fearful of the viruses and radioactive isotopes in the lab, they didn’t search it as thoroughly as they might have. Otherwise, they might have found sensitive documents that Monod would hide inside the leg of a mounted giraffe outside his office. In any case, the Germans soon routed the short-lived Musée de l’Homme group

Monod’s wife, Odette, was the granddaughter of Zadoc Kahn, the former chief rabbi of France. Since the Vichy government soon began enacting Nazi policies, including anti-Jewish laws, and because of homegrown French anti-Semitism, Odette sought refuge for herself, and for her and Jacque’s twin sons (born in August 1939, four weeks before the war broke out), under assumed names, in a village outside of Paris. Meanwhile, Jacques had to register with the Vichy authorities as the spouse of a Jewish person.

With Odette and the children concealed, Monod joined the most militant unit in the Resistance; the Communist-led Franc-Tireurs (Free Shooters) group. Monod was not then a Communist Party member. Nonetheless, he joined the Franc-Tireurs since they actually were fighting the Germans—assassinating German officers in the streets and carrying out sabotage. One of his missions for the Franc-Tireurs took him to Geneva—through the Alps to avoid arrest—to request money for arms from the United States Office of Strategic Services; the precursor of the present Central Intelligence Agency.

By this time, Monod had gone completely underground. He wore a disguise during the day, slept in safe houses at night, and stayed away from his laboratory at the Sorbonne. But then, Andre Lwoff, the head of microbial physiology at the Pasteur Institute, offered Monod a refuge and a place to work in his laboratory at the Pasteur Institute. Monod then led a double-life. By day, as Monod, he worked on his experiments at the Pasteur Institute. At night, he carried out his duties for the Franc-Tireurs, as “Marchal” (from a character in a novel by Stendhal), and as  commander “Malivert.” [Lwoff too had been active in the Resistance, gathering intelligence for the Allies, while also hiding downed American airmen in his apartment.]

Jacques Monod’s identity card for the French Forces of the Interior (FFI), in his nom de guerre ‘Malivert.’

Monod was resolutely committed to the Resistance, while also maintaining a productive research program. At the Pasteur Institute, he and his student, Alice Audureau, made key discoveries that would lead to the later breakthroughs he would make with Jacob. [For instance, Monod and Audureau discovered mutations in E. coli genes that caused the induction of lactose metabolism; a finding that would have important implications concerning gene action and regulation.] Moreover, he was devoted to Odette and their twin sons, and managed to make frequent clandestine visits to see them.

Monod took on increasing responsibilities in the Franc-Tireurs, as more members of the group were discovered and executed by the Germans. In fact, by the time of the allied invasion of Normandy in June 1944, Monod, had become chief of staff of the operations bureau for the National Resistance Organization; a position from which his three predecessors had disappeared (4). As such, Monod prepared battle plans for the allied surge to Paris. He also arranged parachute drops of weapons, railroad bombings, and mail interceptions.

Interestingly, Monod also recruited to the Resistance renowned French chemist, John Frédéric Joliot-Curie (Aside 1), who devised a unique recipe for Molotov cocktails, which were the Resistance’s principal weapon against German tanks. In addition, Monod organized the general strike that facilitated the liberation of Paris. Then, after the liberation of Paris, he became an officer in the Free French Forces, and a member of General de Lattre de Tassigny’s general staff.

[Aside 1: John Frederick Joliet was working as an assistant to Marie Curie, when he married Marie’s daughter, Irene. Afterwards, both John Frederick and Irene changed their surnames to Joliot-Curie. In 1935, the couple was awarded the Nobel Prize in Chemistry for their seminal research on radioactivity. John Frederick then worked at the Collège de France on controlled chain reactions. His work on that was cited by Albert Einstein in his famous 1939 letter to President Franklin Roosevelt, warning Roosevelt of the possibility of a nuclear weapon: “In the course of the last four months it has been made probable through the work of Joliot in France as well as Fermi and Szilard in America—that it may be possible to set up a nuclear chain reaction in a large mass of uranium, by which vast amounts of power and large quantities of new radium-like elements would be generated. Now it appears almost certain that this could be achieved in the immediate future…This new phenomenon would also lead to the construction of bombs…” The Nazi invasion ended Joliet-Curie’s nuclear research. Nevertheless, he managed to smuggle his research notes out of France to England.]

Francois Jacob, a Jewish, nineteen-year old 2nd-year medical student, was planning on a career in surgery when the German occupation of France began in the Spring of 1940. Resolved to carry on the fight against Hitler, Jacob left medical school and boarded one of the last boats for England. In London, he was one of the first of the French to join Charles de Gaulle’s Free French Forces. He wanted to enroll in a combat unit, but, despite his incomplete medical training, he was commissioned as a medical doctor, and then served as a medical officer in North Africa. His surgical career was prematurely cut short in August 1944, when he was severely wounded at Normandy; by a bomb dropped from a German Stuka dive bomber. At the time, he was tending to a dying officer.

For their wartime service, Jacob and Monod were each awarded France’s highest honors for valor. Jacob was awarded the Cross of Liberation, as well as the Légion d’Honneur and the Croix de Guerre. Monod likewise received the Légion d’Honneur and the Croix de Guerre, as well as the American Bronze Star.

Unable to practice surgery after the war because of his wartime wounds, Jacob eventually turned to a career in science. He was accepted at the Pasteur Institute, where he beseeched Lwoff (Monod’s host at the Pasteur Institute) to serve as his mentor. Lwoff rebuffed Jacob several times, but finally agreed to take the young doctor under his wing. Then, in the cramped quarters of Lwoff’s laboratory at the Pasteur, Jacob and Lwoff’s student, Elie Wollman, began a fruitful collaboration that produced key insights into bacterial conjugation and the regulation of lysogeny (Aside 2). After that, Jacob and Monod forged their extraordinary collaboration that would lead to their Nobel Prizes. Note that Jacob’s earlier work with Wollman, on lysogenic induction, would provide the underpinning for his later work on gene regulation with Monod (3).

[Aside 2: Elie Wollman, born in 1917, was Jewish. In 1940, he escaped from the Nazis in Paris and then worked underground in the Resistance as a physician. His parents, Eugene and Elizabeth Wollman, were Pasteur Institute scientists who were seized by the Nazis in 1943 and sent to Auschwitz. They were never heard from again (3).]

In December of 1939, our other main protagonist, twenty-six-year-old Albert Camus, was an unknown, aspiring writer, working as a reporter and editor for a newly founded left-wing newspaper, Alger Republican, in his native Algeria; which was then under French control. Camus was completely opposed to the war, which he saw as “another unnecessary, avoidable, disastrous, absurd chapter of history that would consume the lives of those who did not make it or wish for it.” His antiwar editorials in the Alger Republican outraged French government officials who were calling for unity against Germany. The government finally shut down the newspaper, leaving Camus unemployed. So, Camus returned to France, where the prospects for employment were now better because wartime mobilizations had left many businesses shorthanded. See Aside 3.

[Aside 3: Camus started writing The Stranger while in Algeria, basing it on people and places he knew there. His purpose in The Stranger was to express how one might react to his philosophical notion of the “absurd”—the disconnect between our desire for a rational existence, and the actual world, which appears confused and irrational—in the form of a novel. Meursault, the narrator, and principle character in The Stranger, shows no grief over his mother’s death, no remorse over having committed an unintended murder, and no belief or interest in god. Even while Meursault was awaiting the guillotine, he was reconciled to “the tender indifference of the world.” Meursault’s honesty in describing his feelings makes him a ‘stranger’ in the setting of the novel, and seals his fate.]

Camus was not called up for military service when he returned to France, because he had contracted tuberculosis in Algeria, when he was 17 (Aside 4). Nonetheless, he twice attempted to enlist—the second time when the French Army was on the verge of surrender to the Nazis—to express his solidarity with those who were being drafted. In any case, the military rejected him each time because of his tuberculosis. So, he managed to get a job in Paris as a layout designer for the newspaper Paris-Soir.

[Aside 4: In the pre-antibiotic era, tuberculosis was often fatal, and the 17-year-old Camus indeed had a close brush with death. That experience had a profound effect on the “precocious philosopher,” who made notes on the question of “how, in the light of the certainty of death, one should live life.”]

Parisians began fleeing from their city when the German invasion began in May of 1940. Then, in June, as the Germans were on the verge of entering Paris, the stream of refugees became a flood, with about 70 percent of the city’s metropolitan population of nearly five million eventually taking flight from the city. All Parisian newspapers stopped publishing. However, Paris-Soir hoped to resume its operations in the south, with a reduced staff. Thus, Camus joined the stream of refugees, driving an automobile (almost all the paper’s regular drivers had been drafted), with a Paris-Soir executive as his passenger. After Camus and his passenger were well on their way, Camus suddenly realized that in the rush to vacate from Paris, he may have left his manuscript for The Stranger behind in his room. “He jumped out of the car and threw open the trunk, and was relieved to find in his valise the complete text of The Stranger.” See Aside 5.

[Aside 5: In 1885, Joseph Meister, at nine-years-of-age, was the first recipient of Louis Pasteur’s rabies vaccine and, as an adult, was caretaker of the Pasteur Institute; a position that he still held at the start of the Nazi occupation in 1940.  In despair over the fall of France, and wrongly believing that German bombs killed his family after he sent them away, he went to his apartment, closed the windows, and turned on the gas in his stove (5, 6).]

Camus went with Paris-Soir to Clermont-Ferrand. There, the paper began to publish again, using printing facilities made available by Pierre Laval, the former premier, and now architect of the Petain Vichy government. But with the paper now under Laval’s control, it began publishing anti-Semitic articles, and other articles in support of the Vichy government. Camus did not write any of these items. In any case, he was let go by Paris-Soir after the draftees of the 1940s were discharged and could return to work. Camus then went back to Algeria, where he completed The Stranger.

In 1942, with The Stranger about to be published in France, Camus suffered a nearly fatal relapse of his tuberculosis. He wanted to return to France for treatment in the Massif Central mountain range, but several months would pass before Algerian authorities gave him permission to do so. Then, upon returning to Paris, he would have a purpose that would totally engage him.

One night, under an assumed name (because of the need for secrecy in the Resistance), Camus stole into the clandestine headquarters of Combat (the journalistic arm and voice of the French Resistance), to implore the staff to take him on since he “had already done a little journalism” and would be happy to help in any way. Like Monod, Camus then led a double-life, carrying out his duties at Combat, as “Bauchard.” At first, he helped to select and edit articles, and prepare the paper’s layout. Then, in 1943 he became the paper’s editor, and wrote stirring editorials, exhorting Frenchmen to act against the German occupiers. By the time The Stranger was published in 1942, his recognition as Camus led to his acceptance into the literary and artistic circle that included Sartre, Simone de Beauvier, and Picasso.

Albert Camus’s false identity card, in the name of Albert Mathé, writer. All of the information on the card — birthdate, place, parents — is false.

Camus was suffering from recurrent bouts of tuberculosis all the while that he was carrying out his work at Combat. Nonetheless, as Camus, he also published his essay, The Myth of Sisyphus, which, like The Stranger, contemplates the experience of the Absurd (see Aside 3, above). And he also wrote The Plague, which depicts a city’s response to an outbreak of bubonic plague; perhaps a metaphor for the Nazi occupation. Remarkably, no one at Combat had an inkling that the man who at first had been editing and arranging pages for them as Bauchard was in fact the now renowned Camus. See Aside 6.

[Aside 6: Among laypeople, Jacques Monod is perhaps best known for his “popular” book, Chance and Necessity, published in 1970, and a bestseller in its day. Monod’s Chance and Necessity, and Camus’ The Myth of Sisyphus, are each relevant here because they point up how Camus influenced Monod’s view of the meaning of life. While Camus took a philosophical approach to that issue, Monod’s assessment was also informed by his knowledge of life’s fundamental molecular mechanisms. With the 1953 discovery by Watson and Crick of the molecular structure of DNA, it was apparent how accidental, random, unpredictable mutations in the sequence of bases in DNA were the source of all biological diversity. Thus, Monod knew that all living forms, including humans, are the products of chance genetic mutations and circumstances: “Man at last knows that he is alone in the unfeeling immensity of the universe, out of which he emerged only by chance. Neither his destiny nor his duty have been written down. The kingdom above or the darkness below: it is for him to choose.” [Monod’s title, Chance and Necessity, is from Democritus’ dictum “Everything in the universe is the fruit of chance of chance and necessity.”]  

That we live in a world that is indifferent to our hopes and suffering was the reason for Monod to inquire into the meaning of life, which, for Camus, was “the most urgent of questions.” Camus was often branded an existentialist, but unlike many contemporary existentialist thinkers, Camus vehemently rejected nihilism. In The Myth of Sisyphus, he wrote that Sisyphus gave his life meaning by choosing to believe that he remained the master of his own fate, even though he was condemned to rolling his rock uphill each day, only to have it roll back down.

On the opening page of Chance and Necessity, Monod includes a lengthy quotation from the closing paragraphs of The Myth of Sisyphus. “The struggle itself towards the heights is enough to fill a man’s heart…One must imagine Sisyphus happy.” Camus is advocating that we oppose the certainty of death in an uncaring Universe by living life to the fullest. For Monod, life is like Sisyphus, pushing its rock uphill. The end might be bleak, but “the struggle towards the heights is enough to fill a man’s heart.”]

By 1944, the liberation of Paris was imminent, Combat went from a monthly publication to a daily one, and the paper chanced to circulate in the open. Camus was still writing his editorials anonymously. And when his identity was finally revealed, his inspiring, eloquent words resulted in his widespread public acclaim.

Monod and Camus were very likely aware of each other at this point in our saga, but they had not yet met. Their meeting would happen after the liberation of France, and it would be in response to a new totalitarian threat; from the Soviet Union. It transpired as follows.

In 1948, Monod was working full-time on his research at the Pasteur Institute, when events in the Soviet Union moved him to write a stirring editorial that appeared on the front page of Combat. [Camus had left Combat the previous year, after it became a commercial paper.] Monod’s piece was in response to a pseudoscientific doctrine advanced by Stalin’s head of Soviet agriculture, Trofim Lysenko, which asserted that organisms could swiftly change their genetic endowment in response to a new environment. [Lysenko’s doctrine is reminiscent of discredited Lamarckian doctrine, also known as heritability of acquired characteristics—i.e., the premise that if an organism changes to adapt to an environment, it can pass on those changes to its offspring.] Lysenko based his doctrine on his purported discovery of a means to enable winter wheat to be sown in the spring.

Stalin embraced Lysenkoism—during an acute grain shortage in Russia—since it was in accord with his ideology to create the New Soviet Man. Stalin also banned all dissent against Lysenko’s doctrine. Consequently, traditional Russian geneticists were exiled or murdered, Mendelian genetics was no longer practiced in the Soviet Union, and Soviet agriculture suffered severely.

Monod was roused to write his editorial after French Communist newspapers began to widely disseminate Lysenko’s doctrine in France. One Party newspaper proclaimed Lysenko’s discovery “A Great Scientific Event,” and further asserted that the notion of evolution by natural selection was a racist form of thinking, in harmony with Nazi doctrine (7). Another Party newspaper condemned Mendelian genetics for being “bourgeois, metaphysical and reactionary,” while claiming that it must be false because it is reactionary; having been invented by an Austrian monk. In Contrast, Lysenkoism is true because it is progressive and proletarian.

A Party member’s position on Lysenko indeed had become a gauge of his commitment to Stalin’s Soviet cause. But for Monod, the Soviet embrace of Lysenko was “senseless, monstrous, unbelievable.” As expected, Monod’s article was strongly condemned by the powerful French Communist Party, which enjoyed broad support from both intellectuals and workers; many of whom saw the Soviet Union as a model for a French socialist state. In any case, the Party’s strong backlash inspired Monod to “make his life’s goal a crusade against anti-scientific, religious metaphysics, whether it be from Church or State.” Importantly, a separate consequence of the Lysenko affair was that it influenced François Jacob to focus his research in the field of genetics. See Aside 7.

[Aside 7: Ironically, the observation that Jacob and Monod initially set out to explain looked remarkably like Lysenkoism. When E. coli are fed a solution of glucose and lactose, they grow rapidly until glucose—their preferred carbon source—is depleted. Only then, they turn to metabolizing lactose. But, in contrast to Lysenko’s doctrine, Jacob and Monod showed that when E. coli “adapts” to lactose, it does so without changing its genes. Instead, the genes encoding the enzymes that metabolize lactose lie dormant until lactose induces them, under conditions in which glucose is not available. That is, Jacob and Monod determined that lactose regulates lactose metabolism in the cell by acting as an inducer of genes that already exist in the cell; as opposed to lactose causing the cell to undergo a Lamarckian acquisition of a genetic characteristic. In so doing, Jacob and Monod created the now well-established paradigm of inducers, regulators, regulator genes, and operators.]

While Monod was crusading against Lysenkoism, Camus was having his own feud, in public, with Sartre, who had chastised him for his anti-Soviet stance. Camus had once been a Communist, in Algeria, mainly because he was troubled by the way in which the European French treated the native Algerians. However, he was never very sympathetic to the Marxist cause. Monod too had once been a member of the Communist Party; but only because it enabled him to have a voice in the running of the Resistance. In any case, Camus seized upon Monod’s condemnation of Lysenkoism in his feud with Sartre.

Our two main protagonists finally met when Camus co-founded the anti-Stalin, anti-totalitarian Groupes de Liaison Internationale. Monod attended one of the group’s meetings. There, he, and Camus, discovering that they shared much in common, forged their friendship. Carroll writes: “Camus, who so treasured the sense of solidarity that existed among the Resistance, had in Monod a new comrade who shared both the deep bond of that wartime experience and an unqualified opposition to a new common enemy.”

As noted, Monod’s views on the meaning of life owed much to Camus. Likewise, Camus learned from Monod. Camus not only used Monod’s case against Lysenko in his dispute with Sartre, but he also “borrowed” from Monod in The Rebel; in which Camus argued that revolution inevitably leads to tyranny. In any event, after Camus and Monod had separately fought the Nazis, they were now united against another oppressor—the totalitarian state run by Stalin. [Camus’ anti-Soviet stance cost him the friendships of many French intellectuals on the left. He and Sartre never spoke to each other again.]

Monod was also troubled by the situation of scientists working under Eastern European Soviet regimes. In 1959, he organized the escape into Austria of Hungarian biochemist Agnes Ullman (who participated in the failed Hungarian uprising of 1956), and her husband, also a scientist. Earlier, in 1958, Agnes Ullman managed to visit Monod at the Pasteur Institute, and confided to him that she and her husband wanted to defect from Hungary. Monod maintained contact with the Ullmans in Hungary, using coded messages, written in invisible ink, which turned blue when exposed to iodine. The Ullmans finally crossed into Austria, hidden underneath a bathtub, in a compartment of a pull-along camping trailer. See Aside 8.

[Aside 8. Agnes Ullmann, became Monod’s long-time close collaborator at the Pasteur Institute. Now retired, she was carrying out research at the Pasteur Institute as recently as 2012; 53 years after her rescue from Hungary. At the Institute, she collaborated with Monod on characterizing the lac operon promoter, on complementation between β-galactosidase subunits, and on the role of cAMP in overcoming the repressive effect of glucose (catabolite repression) on lactose metabolism in E. coli.]

There are numerous other instances in which Monod stepped forward to fight injustice and defend human rights. In 1952, he wrote a letter in Science that might have been “ripped from today’s headlines.” It protested the U.S. government’s rejection of visa requests for himself and other Europeans who had once been Communists. Monod also condemned the treatment of Jews in the Soviet Union, while continuing to speak out against Soviet totalitarianism in general. And, in 1965, shortly after Monod, Lwoff, and Jacob received word of their Nobel Prizes, they publicly appealed to the French government to allow the use of contraceptives, and the legalization of abortion. See Aside 9.

[Aside 9: Jacob too was devoted to the defense of human rights. He chaired a committee of the French Academy of Sciences that supported persecuted scientists living under totalitarian regimes, and he worked for the release of those who had been imprisoned for their political views. Moreover, he forcefully advocated for the public support of the biological and medical sciences. What’s more, Jacob also had a distinguished writing career that produced a series of acclaimed books, including The Logic of Life: A History of Heredity; Of Flies, Mice and Men; The Possible and the Actual, and his memoir, The Statue Within. In Joshua Lederberg’s review of the latter for The Scientist, he stated:As a work of literature, it evokes unmistakable overtones of Rousseau, Proust, and Sartre.” In Jon Beckwith’s view, all of Jacob’s books are “written in a fluid and elegant style” Others refer to the “clarity and grace” of Jacob’s writing. See reference 8 for more on Jacob.]

In 1966, Martin Luther King Jr. and Harry Belafonte visited France to raise funds for the Southern Christian Leadership Conference (SCLC). Remarkably, Monod was chosen to introduce King to a crowd of 5,000 people at Paris’ Palais des Sports. Belafonte was introduced by French singer and actor Yves Montand (9).

Coretta Scott King shaking hands with Jacques Monod, as Martin Luther King Jr. looks on. From l-r: French actress Simone Signoret, Harry Belafonte, French actor and singer Yves Montand. 29 March 1966, at a meeting of the “Movement for the Peace.”

The intellectual lives of Monod and Camus played out in entirely different areas. Yet the parallels were striking. Each, in his way, searched for meaning in life. Moreover, each put his life on the line to oppose ignorance, injustice, and totalitarianism. And, it is clear from their correspondences that they were dear to each other. Here is the note from Monod that elicited Camus’ response at the top of this post.

My dear Camus,

My emotion and my joy are profound. There were many times when I felt like thanking you for your friendship, for what you are, for what you managed to express with such purity and strength, and that I had likewise experienced. I wish that this dazzling honor would also appear to you, in some small part, as a token of friendship and of personal, intimate recognition. I would not dare coming to see you right now, but I embrace you fraternally.

Jacques Monod

This piece ends with a few personal thoughts. Jacques Monod was a Nobel Prize-winning scientist, a hero of the French Resistance, a rescuer of persecuted scientists from behind the Iron Curtain, and a leading voice against tyranny and oppression. And, he was also blessed with dashing good looks. I remember well the women among my fellow graduate students in the 1960s finding him to be very attractive. But, on a more serious note: Today, when political and religious blocs dismiss evidence-based science in favor of alternative ‘facts’ in order to advance their ideologies, and when they are tacitly aided by a press that all too often gives equal validity to all points of view, and while scientists seem to be groping for an effective response, one can hope that scientists with the courage, eloquence, and eminence of Jacques Monod and Francois Jacob might emerge to take up the cause of science and reason. Meanwhile, it is especially important for young scientists, and the public, to be aware of the examples set by these men. See Aside 10.

[Aside 10: The following is from a March 8, 2017 editorial in Nature.  “Last week, state legislators in Iowa introduced a bill that would require teachers in state public schools to include ‘opposing points of view or beliefs’ in lessons on topics including global warming, evolution and the origins of life… Since last month, Indiana, Idaho, Alabama, Texas, Oklahoma and Florida have all introduced and discussed similar tweaks to the way in which they want to educate their children… Although these proposed changes are typically presented by their supporters as giving teachers the chance to discuss genuine scientific controversies, in truth they are (very) thinly veiled attempts to pursue political and religious agendas that have no place in school science lessons — for whatever age. They seek to import the alternative facts and misleading rhetoric of the new federal government and to impose it on children who deserve much better from those elected to serve them.”]


  1. A Most “Elegant” Experiment: Sydney Brenner, Francois Jacob, Mathew Meselson, and the Discovery of Messenger RNA , Posted on the blog October 6, 2016.
  2. Sean B. Carroll, Brave Genius: A Scientist, a Philosopher, and their Daring Adventures from the French Resistance to the Nobel Prize, Crown, 2013.
  3. Genealogies and a Selective History of Lysogeny: Featuring Friedrich Loeffler, Emile Roux, Andre Lwoff, Elie Wollman, and Francois Jacob, Posted on the blog January 28, 2015 (8).
  4. Agnes Ullmann, In Memoriam: Jacques Monod (1910–1976), Genome Biology and Evolution 3:1025-1033. 2011, DOI: https://doi.org/10.1093/gbe/evr024.
  5. Louis Pasteur: One Step Away from Discovering Viruses, Posted on the blog January 7, 2015.
  6. Dufour, H. D., and S. B. Carroll, History: Great myths die hard, Nature 502:32–33, 2013.
  7. John Marks, Jacques Monod, François Jacob, and the Lysenko Affair: Boundary Work, L’Esprit Créateur: Genetics and French Culture. 52:75-88.
  8. Beckwith, J., and M. Yaniv, Francois Jacob (1920-2013), Current Biology 23:R422-R425, 2013.
  9. Remembering Dr. Martin Luther King Jr., CBS Minnesota, January 19, 2013, 3:51 PM.