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History of the Discovery of Hepatitis a Virus Downloaded from http://perspectivesinmedicine.cshlp.org/ on October 1, 2021 - Published by Cold Spring Harbor Laboratory Press History of the Discovery of Hepatitis A Virus Stephen M. Feinstone Department of Biochemistry and Molecular Medicine, George Washington University School of Medicine, Washington, D.C. 20037 Correspondence: [email protected] Disease outbreaks resembling hepatitis A have been known since antiquity. However, it was not until World War II when two forms of viral hepatitis were clearly differentiated. After the discovery of Australia antigen and its association with hepatitis B, similar method- ologies were used to find the hepatitis A virus. The virus was ultimately identified when investigators changed the focus of their search from serum to feces and applied appropriate technology. iseases resembling hepatitis A, both in DIFFERENTIATION OF TWO FORMS OF Dindividuals and in outbreaks involving VIRAL HEPATITIS groups, were reported in China as early as 5000 years ago. Hippocrates noted a disease Viral hepatitis was a major problem for both the he called benign epidemic jaundice in “De Allies and the Axis during World War II. Early Morbis Internis” that certainly resembled hep- in the war, an outbreak of hepatitis related to atitis A. More accurate descriptions of hepatitis yellow fever vaccine, stabilized with human se- A began appearing in the 17th century often rum involving 49,233 clinically apparent cases associated with military campaigns. The first (Seeff et al. 1987), prompted a major hepatitis outbreak recorded in the United States was research effort. As the records on the vaccinees in 1812 in Norfolk, VA, and the disease was were very good, the incubation period was de- common among the Union troops during fined accurately as between 60 and 154 days. www.perspectivesinmedicine.org the Civil War with more than 40,000 cases re- This outbreak as well as the more widespread ported. The association of hepatitis A with war problem of infectious hepatitis caused both the led to 19th-century terms such as “kriegsik- British and the Americans to initiate studies of terus” or “jaunisse des camps.” Hepatitis A viral hepatitis. It became clear that one form continued to afflict troops on both sides during of hepatitis spread quickly among troops and World War I and in the Second World War by 1943 hepatitis had become a real hindrance when there were estimates of 16 million cases of the war effort in North Africa and the of hepatitis among combatants and civilians Mediterranean. U.S. Army epidemiologic stud- (Sherlock 1984; Gust and Feinstone 1988; Fon- ies showed that epidemic or infectious hepatitis seca 2010). had a much shorter incubation period than se- Editors: Stanley M. Lemon and Christopher Walker Additional Perspectives on Enteric Hepatitis Viruses available at www.perspectivesinmedicine.org Copyright © 2018 Cold Spring Harbor Laboratory Press; all rights reserved Advanced Online Article. Cite this article as Cold Spring Harb Perspect Med doi: 10.1101/cshperspect.a031740 1 Downloaded from http://perspectivesinmedicine.cshlp.org/ on October 1, 2021 - Published by Cold Spring Harbor Laboratory Press S.M. Feinstone rum hepatitis—18 to 25 days compared with a infections of some children and were controver- mean of 90 days for hepatitis following the yel- sial even during the era in which they were low fever vaccine (Havens 1968). Second, they conducted. They were justified by the investiga- found that some soldiers that had developed tors based on their belief that every child who hepatitis after immunization with the yellow entered the facility developed hepatitis soon fever vaccine could still develop a second bout after arrival. In the course of these studies, the of infectious hepatitis. They also noted that investigators identified a child with two distinct officers were more likely to develop infectious bouts of hepatitis from whom serum samples hepatitis than enlisted men and that the disease had been collected during the acute and recov- often followed outbreaks of diarrheal disease, ery stages of each bout. Additional studies suggesting a fecal–oral mechanism of spread. with these two samples, termed MS-1 for short These epidemiologic studies were accompa- incubation (infectious) hepatitis and MS-2 for nied by experimental infections of humans per- long incubation (serum) hepatitis, further de- formed by the Americans and British as well fined two distinct hepatitis viruses and, most as the Germans (Voegt 1942; Neefe et al. 1944, importantly, established a well-characterized, 1946; MacCallum et al. 1951). Transmission to known infectious inoculum for each (Krugman volunteers was not always successful. Preexisting et al. 1967). Krugman also determined the immunity in a high proportion of volunteers, as general period of infectivity for persons infected well as the inability to know whether any given with these viruses, showed that serum hepatitis inoculum was actually infectious, caused the often became chronic, and that standard results of these studies to be difficult to interpret. pooled human immune serum globulin was Nevertheless, these studies pointed to two protective against infectious hepatitis (Krugman distinct diseases, one with a short incubation and Giles 1970, 1972). This latter finding con- period transmitted by the fecal–oral route and firmed earlier studies done by the U.S. Army the other transmitted by serum with a relatively during World War II (Gellis et al. 1945). long incubation period. The investigators also Because of the continuing threat to military showed a lack of cross immunity between the operations posed by infectious hepatitis, the U.S. two types of infections and that some of the Army continued to sponsor research after the physical characteristics of the causative agents war under the aegis of the Armed Forces Epide- were distinct. These two forms of hepatitis miological Board. To obtain larger quantities became known generally as “infectious hepati- of the infectious hepatitis inoculum, the army tis” and “serum hepatitis.” It was not until the contracted for studies in which the Krugman early 1950s that the first description of these MS-1 inoculum was amplified by infecting www.perspectivesinmedicine.org diseases as “type A” and “type B” viral hepatitis volunteer inmates at the federal prison in Joliet, appeared in a report from an expert committee IL (Boggs et al. 1970; Melnick and Boggs 1972). of the World Health Organization (MacCallum Volunteers were inoculated with the MS-1 strain 1953). of hepatitis A, and the investigators collected Following the war, virology entered its gold- stools and serum samples throughout the course en era with the advent of tissue culture made of the illness, from the incubation period to possible by the development of defined media acute disease, and then during convalescence. and antibiotics (Robbins and Enders 1950). In addition, MS-1 infection was passed from Many viral agents were identified during this one volunteer to another. period but neither hepatitis agent was success- fully propagated in cell culture. Saul Krugman, EARLY ATTEMPTS TO ISOLATE AN Robert Ward, and Joan Giles conducted studies INFECTIOUS HEPATITIS A AGENT between 1956 and the early 1970s at the Willow- brook State School for intellectually handi- In early quests to isolate an infectious agent, capped children on Staten Island, New York. classic virologic methods, including inocula- These studies involved deliberate experimental tions of laboratory animals and embryonated 2 Advanced Online Article. Cite this article as Cold Spring Harb Perspect Med doi: 10.1101/cshperspect.a031740 Downloaded from http://perspectivesinmedicine.cshlp.org/ on October 1, 2021 - Published by Cold Spring Harbor Laboratory Press History of the Discovery of Hepatitis A Virus chicken eggs with hepatitis-related material, did Friedrich Deinhardt working at the Presbyteri- not yield positive results. With the development an-St. Lukes Hospital in Chicago, IL, reported of cell culture as a routine laboratory technolo- that a human hepatitis virus could be passaged gy, scientists could regularly propagate mamma- in marmoset monkeys (Saguinus sp., New lian cells in vitro and identify many viruses World nonhuman primates now generally known to be associated with human diseases considered to be tamarins) and that infected (Robbins and Enders 1950). animals developed evidence of liver disease Various cell culture methods were applied to (Deinhardt et al. 1967). One hepatitis-inducing the identification of the causative agent of hep- inoculum, GB, came from a surgeon who had atitis A (MacCallum 1972), but the most exten- developed hepatitis without a known exposure. sively reported study was performed by Wilton Deinhardt did extensive work to characterize the Rightsel and his colleagues at the Parke-Davis GB agent, but he could not specifically associate Company using a cell line termed Detroit-6, it with hepatitis A. Wade Parks and Joseph Mel- which was derived from human bone marrow nick, at Baylor College of Medicine in Houston, (Rightsel et al. 1956, 1961). These investigators TX, determined that the GB agent was likely a identified a cytopathic effect (CPE) in cells marmoset virus, but they also were unable to inoculated with various specimens thought to prove that these primates could be infected contain infectious virus. This CPE could be with the MS-1 virus (Parks and Melnick 1969; neutralized by convalescent serum from pa- Parks et al. 1969). The story of GB resurfaced tients with hepatitis A–like illnesses. Although in 1995 when two viruses, GBV-A and GBV-B, the neutralization studies were presumptive were identified by molecular cloning in animals evidence for specificity, further studies were in- infected with the eleventh animal passage of consistent for both the CPE and neutralization. the GB agent (Simons et al. 1995). Neither of The Detroit-6 cell culture studies were eventu- these agents infect humans. A related virus, ally abandoned at Parke-Davis, but Ruth Cole, a GBV-C, has been identified in humans, but member of the Parke-Davis team, returned to it is of very low or nonexistent pathogenicity.
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