Otto Loewi was the son of the Jewish wine merchant Jakob Loewi and his wife Anna, née Willstädter. He attended the Woehlerschule (elementary school) in Frankfurt am Main from 1879 and transferred to the humanistic Goethe Gymnasium in 1882, where he graduated on September 4, 1891.1
Loewi began studying medicine at the Kaiser Wilhelm University in Strasbourg that same year at his parents' request, where he initially attended lectures primarily in history of art, architecture, and theoretical philosophy. After completing his initial medical exams, he moved to Munich for the winter semester of 1893/94. He returned to Strasbourg one year later. As he was fascinated by the lectures and courses held by Bernhard Naunyn (1839-1925), Loewi decided to pursue a career in clinical medicine. He passed the state examination in 1896 and was awarded a doctorate in medicine under Oswald Schmiedeberg (1838-1921) the same year, with an experimental pharmacological thesis on the effects of hydrocyanic acid, arsenic and phosphorus on isolated frog hearts.2 In the course of his doctoral work, he met scientists such as Oskar Minkowski (1858-1931), Walter Straub (1874-1944), Arthur Cushny (1866-1926), Karl Spiro (1896-1932), and Franz Hofmeister (1850-1922), through whom his interest in pharmacological and biochemical questions was encouraged and fostered. Loewi began studying chemistry under Martin Freund (1863-1920) at the Biochemical Institute of the University of Frankfurt in 1896. He continued his studies under Hofmeister in Strasbourg.
University Career
Loewi took a position as a resident in internal medicine under Carl von Noorden (1858-1944), a specialist in metabolic diseases and nutritional therapy, at the Städtisches Krankenhaus in Frankfurt am Main in 1897. Based on his experience there, he devote himself to basic medical research, or rather clinical pharmacology and metabolic research instead of clinical work. He moved to the Philipps University in Marburg after just one year, where he became a research assistant to Hans Horst Meyer (1853-1939), the director of the Pharmacological Institute, where he habilitated on November 19, 1900.3
Loewi spent several months at the renowned institute of physiologist Ernest Starling (1866-1927) in London in 190, to further his education in scientific methodology and laboratory techniques and to establish contacts in British science. This is also where he met Henry Hallet Dale (1875-1968), with whom he would later receive the Nobel Prize, and would maintain close scientific and friendly contact.
Loewi was appointed associate professor in Marburg in 1904, and was appointed acting director of the Pharmacological Institute on November 1. After Meyer had been appointed to Vienna, Loewi moved there too one year later. He primarily worked with Alfred Froehlich (1871-1953) and began research on the nervous system during this time. Loewi finally accepted an appointment as full professor of pharmacology at the Karl Franzens University in Graz in 1909. He remained in this chair until 1938, and turned down numerous offers from other universities.
Loewi had assumed Austrian citizenship in addition to German citizenship in the meantime. He married Guida Henrietta Goldschmiedt (1889-1958), the daughter of Guido Goldschmiedt (1850-1915) in 1908. He had met Goldschmiedt – a full professor of chemistry working in Prague and Vienna – during his annual vacation in Pontresina, Switzerland, the year before. The couple had three sons and one daughter, Hans (*1909), Anna (*1911), Viktor (*1913) and Guido (*1915). The family acknowledged their Jewish identity, but were not religious and were considered assimilated.4
The DGIM, chaired by Paul Morawitz (1879-1936), and the German Pharmacological Society, chaired by Otto Loewi, decided to hold their annual congresses jointly in Wiesbaden in 1932. The joint congress coincided with the 50th anniversary of the founding of the Society for Internal Medicine. Franz Volhard (1872-1950), and Sir Henry Dale from London were the main speakers and the first key topic to be addressed was the circulatory effects of endogenous substances.5
Scientific Discoveries and Nobel Prize 1936
Loewi's research interests were broad. He first gained international recognition with a study published in 1902, in which he demonstrated the body's ability to build proteins from amino acids.6 He thus laid important foundations for the nutritional work of Frederick G. Hopkins (1861-1947), who had discovered vitamins and had been awarded the 1929 Nobel Prize for Medicine.
Loewi's research interests were broad.
With the help of experiments conducted on frog hearts, Loewi, in the spring of 1921, was able to demonstrate that the transmission of nerve impulses to (heart) muscle(s) is chemical.7 To do this, he stimulated the vagus nerve of a frog heart soaked in saline and transferred the saline to a denervated frog heart, which then slowed down, corresponding to a vagus response. This confirmed the theoretically established but hitherto controversial research hypothesis of chemical transmission of nerve impulses to muscles.
According to his own statements, the idea for the experiment had come to Loewi in a dream on Easter 1921 and had tempted him, still half asleep, to implement it directly in the early hours of the morning.8 The chemical transmitter substance he initially called "vagus substance" was identified by Henry Dale as acetylcholine. Thus the first neurotransmitter had been found and identified. Both researchers were honored for their discoveries on the chemical transmission of nerve impulses with the Nobel Prize in Physiology or Medicine in 1936.
Persecution, Expropriation and Forced Exile
With the "Anschluss" of Austria to the German Nazi state, Loewi and his two younger sons were taken into "protective custody" in Graz for nearly two months on March 12, 1938, along with numerous other Jewish citizens. His release was conditional on his undertaking to leave the country voluntarily within two months.9 Loewi's disenfranchisement and imprisonment quickly became known in his professional circle. His colleagues from abroad, especially from Great Britain and Sweden, sent numerous letters to Graz to obtain Loewi's release.10 The Berlin pharmacologist Wolfgang Heubner (1877-1857) spoke publicly as the host of the 14th meeting of the German Pharmacological Society at the Langenbeck-Virchow House on April 24, 1938 in a well-received speech: "Many souls are touched, when misfortune befalls an outstanding pioneer of far-reaching correlations."11
The University of Graz put Loewi into provisional retirement at the end of May 1938. Innsbruck full professor Adolf Jarisch (1891-1965) was to be appointed to his chair as his successor, but he declined.12 Loewi asked for permission to emigrate and was allowed to leave a few months later on the condition that he not engage in any scientific activity abroad. Furthermore, the Gestapo forced him to transfer his Nobel prize award money to a Nazi-controlled bank. All of his real estate was expropriated.13
Loewi fled to England on September 28, 1938, and stayed with Henry Dale for a few weeks until, at the invitation of the Belgian Fondation Francqui, he was offered a visiting professorship at the Institute of Pharmacology under Leo Zunz at the Université Libre in Brussels. He stayed for only one semester, though, as he was unable to return to Belgium, after a vacation in England the following year, as World War II had broken out in the meantime. He accepted a research position with James Andrew Gunn (1882-1958) at the Nuffield Institute for Medical Research in Oxford instead.
Loewi was offered a tenured research professorship at the Pharmacology Department of New York University College of Medicine under George B. Wallace (1874-1948) in 1940. He accepted and arrived in New York on June 1, 1940, two days before his 67th birthday. He was joined by his wife Guida in early 1941, who had not been allowed to leave Austria until that time. She was only permitted to leave, once the compulsory expropriation proceedings had been completed, as a "destitute Jew". The Loewis' children had already emigrated earlier on; their daughter Anna had emigrated to the USA via detours with her husband, the chemist Ulrich Weiss (1908-1989), their son Viktor to Argentina, their son Guido to Canada, and their son Hans first to Peru and then also to the USA.14
"New Home" America
Loewi was granted American citizenship on April 1, 1946. He was a resident of New York (at least temporarily, at 155 East 93 Street) until his death in 1961. Numerous guest lectures took him to various parts of the USA. Woods Hole in Massachusetts became a special home for him, where he spent the annual summer vacation with his wife and enjoyed an intensive exchange with colleagues and students at the Marine Biological Research Station. After his death in New York in 1961, Loewi was buried in Woods Hole in August 1962, in accordance to his wishes.
Loewi was not only very active in the scientific field well into old age, he also remained avidly interested in art (history), music, and theater throughout his life. Artists, musicians and literary figures were also regular guests at the dinner parties the Loewis loved to host.
Memberships, Honors, Obituaries
Loewi was an (honorary) member of numerous scientific societies and academies, including the Physiological Society London, the Deutsche Gesellschaft für Pharmakologie und Physiologie, the Harvey Society New York, and the Società Italiana di Biologia Sperimentale, as well as the Deutsche Akademie der Naturforscher Leopoldina, the Royal Societies London and Edinburgh, and the American Academy for Pharmacology and Experimental Therapeutics.
He had been awarded many honorary doctorates, including by universities such as New York, Yale, Graz, and Frankfurt am Main. He had been permitted to use the title "Hofrat" since 1923. Loewi received many an award, including the "Große Ehrenzeichen für Wissenschaft und Kunst" (Austrian Grand Decoration of Honor for Science and Art) twice (1936, 1959), the Ignaz-Lieben Prize (1924), the Cameron Prize of the University of Edinburgh (1944), the Schmiedeberg Plakette of the German Society for Experimental and Clinical Pharmacology and Toxicology (1957), and the Ring of Honor of the City of Graz (1959).
There were many obituaries. At the 1962 annual congress of the DGIM, of which Loewi had been a member until 1933, conference president Ferdinand Hoff (1896-1988) paid tribute to him, saying, "An inhuman system had also forced this great scholar to emigrate. Not long ago, I was able to spend unforgettable hours with him in New York and to enjoy his sceptical wisdom, in which he acknowledged German science without bitterness. He was one of the greats of medical science.“15
Namesakes
Otto Loewi acts as namesake for, among others, the Forschungszentrum für Gefäßbiologie, Immunologie und Entzündungn in Graz and the Otto Loewi Gesellschaft zur Erforschung der Erkrankungen des autonomen Nervensystems, founded in 2017 and based in Innsbruck. The Neurowissenschaftliche Gesellschaft has been awarding the Otto Loewi Medal every four years to outstanding scientists in the field of neuroscience since 2018.
