{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,7]],"date-time":"2026-07-07T23:26:47Z","timestamp":1783466807921,"version":"3.55.0"},"reference-count":110,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2026,2,9]],"date-time":"2026-02-09T00:00:00Z","timestamp":1770595200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100009619","name":"Research Program on Hepatitis from the Japan Agency for Medical Research and Development, AMED","doi-asserted-by":"publisher","award":["JP25fk0210132"],"award-info":[{"award-number":["JP25fk0210132"]}],"id":[{"id":"10.13039\/100009619","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>Hepatitis E virus (HEV) is an important cause of acute and chronic hepatitis worldwide, transmitted primarily through waterborne exposure and zoonotic foodborne pathways. In recent years, shellfish have attracted growing attention as a potential vehicle for HEV transmission. This interest is driven by epidemiological observations linking shellfish consumption to human HEV infection and by repeated detection of HEV RNA in bivalve mollusks across multiple geographic regions. This review critically evaluates the current evidence by integrating epidemiological data, environmental and food surveillance studies, and mechanistic insights into viral accumulation in shellfish. Signals from outbreak investigations, observational studies, seroepidemiological surveys, and case reports suggest that shellfish may contribute to HEV exposure. However, these findings are largely associative, methodologically heterogeneous, and limited by the absence of explicit documentation of raw or undercooked shellfish consumption in many cases. To date, no study has recovered infectious HEV from shellfish, nor has any established molecular epidemiological linkage between shellfish-derived HEV and human infections. Mechanistic knowledge from norovirus and hepatitis A virus demonstrates that bivalves can bioaccumulate enteric viruses through filter feeding, yet HEV-specific processes governing viral binding, persistence, and infectivity within shellfish remain poorly defined. Surveillance data reveal marked geographic variation in HEV RNA detection among shellfish species and production areas. Overall, existing evidence supports shellfish as a biologically plausible but unconfirmed source of HEV exposure. Addressing key knowledge gaps\u2014particularly through direct infectivity assessments and high-resolution molecular linkage studies\u2014will be essential to determine the public health significance of shellfish within the broader ecology of HEV transmission.<\/jats:p>","DOI":"10.3390\/v18020220","type":"journal-article","created":{"date-parts":[[2026,2,9]],"date-time":"2026-02-09T16:13:32Z","timestamp":1770653612000},"page":"220","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Shellfish as a Potential Source of Hepatitis E Virus: Epidemiological Evidence, Biological Plausibility, and Research Gaps"],"prefix":"10.3390","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0827-0964","authenticated-orcid":false,"given":"Hiroaki","family":"Okamoto","sequence":"first","affiliation":[{"name":"Division of Virology, Department of Infection and Immunity, Jichi Medical University School of Medicine, 3311-1 Yakushiji, Shimotsuke 329-0498, Tochigi, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,2,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1186\/s12985-024-02470-3","article-title":"From discovery to treatment: Tracing the path of hepatitis E virus","volume":"21","author":"Letafati","year":"2024","journal-title":"Virol. J."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1038\/s41575-024-00950-z","article-title":"Hepatitis E virus: From innate sensing to adaptive immune responses","volume":"21","author":"Bruggemann","year":"2024","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1038\/nrgastro.2017.150","article-title":"Hepatitis E virus: Advances and challenges","volume":"15","author":"Nimgaonkar","year":"2018","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"988","DOI":"10.1002\/hep.25505","article-title":"The global burden of hepatitis E virus genotypes 1 and 2 in 2005","volume":"55","author":"Rein","year":"2012","journal-title":"Hepatology"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Primadharsini, P.P., Nagashima, S., and Okamoto, H. (2019). Genetic Variability and Evolution of Hepatitis E Virus. Viruses, 11.","DOI":"10.3390\/v11050456"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1016\/S1473-3099(08)70255-X","article-title":"Hepatitis E: An emerging infection in developed countries","volume":"8","author":"Dalton","year":"2008","journal-title":"Lancet Infect. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1128\/CMR.00057-13","article-title":"Hepatitis E virus infection","volume":"27","author":"Kamar","year":"2014","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1109","DOI":"10.1016\/j.jhep.2022.05.006","article-title":"Chronic hepatitis E: Advancing research and patient care","volume":"77","author":"Ma","year":"2022","journal-title":"J. Hepatol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"482","DOI":"10.4254\/wjh.v14.i3.482","article-title":"Hepatitis E in immunocompromised individuals","volume":"14","author":"Damiris","year":"2022","journal-title":"World J. Hepatol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.clinre.2014.07.005","article-title":"Hepatitis E virus: Chronic infection, extra-hepatic manifestations, and treatment","volume":"39","author":"Kamar","year":"2015","journal-title":"Clin. Res. Hepatol. Gastroenterol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"S196","DOI":"10.3350\/cmh.2024.0758","article-title":"Hepatitis E as a trigger for acute-on-chronic liver failure","volume":"31","author":"Buti","year":"2025","journal-title":"Clin. Mol. Hepatol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1186\/s12985-020-01343-9","article-title":"Hepatitis E virus infection during pregnancy","volume":"17","author":"Wu","year":"2020","journal-title":"Virol. J."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1082","DOI":"10.1016\/j.jhep.2016.11.016","article-title":"Hepatitis E virus: Infection beyond the liver?","volume":"66","author":"Pischke","year":"2017","journal-title":"J. Hepatol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1007\/s00705-008-0045-6","article-title":"Production of monoclonal antibodies against hepatitis E virus capsid protein and evaluation of their neutralizing activity in a cell culture system","volume":"153","author":"Takahashi","year":"2008","journal-title":"Arch. Virol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1112","DOI":"10.1128\/JCM.02002-09","article-title":"Hepatitis E Virus (HEV) Strains in Serum Samples Can Replicate Efficiently in Cultured Cells Despite the Coexistence of HEV Antibodies: Characterization of HEV Virions in Blood Circulation","volume":"48","author":"Takahashi","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"4232","DOI":"10.1128\/JVI.02804-15","article-title":"Distinct Entry Mechanisms for Nonenveloped and Quasi-Enveloped Hepatitis E Viruses","volume":"90","author":"Yin","year":"2016","journal-title":"J. Virol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"e00822-17","DOI":"10.1128\/JVI.00822-17","article-title":"Characterization of the Quasi-Enveloped Hepatitis E Virus Particles Released by the Cellular Exosomal Pathway","volume":"91","author":"Nagashima","year":"2017","journal-title":"J. Virol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"001778","DOI":"10.1099\/jgv.0.001778","article-title":"ICTV Virus Taxonomy Profile: Hepeviridae 2022","volume":"103","author":"Purdy","year":"2022","journal-title":"J. Gen. Virol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1007\/s12560-021-09461-5","article-title":"The Foodborne Transmission of Hepatitis E Virus to Humans","volume":"13","author":"Treagus","year":"2021","journal-title":"Food Environ. Virol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Meng, X.J. (2016). Expanding Host Range and Cross-Species Infection of Hepatitis E Virus. PLoS Pathog., 12.","DOI":"10.1371\/journal.ppat.1005695"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1007\/s12560-015-9207-3","article-title":"Hepatitis A and E Viruses in Wastewaters, in River Waters, and in Bivalve Molluscs in Italy","volume":"7","author":"Iaconelli","year":"2015","journal-title":"Food Environ. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"5870","DOI":"10.1128\/AEM.00336-10","article-title":"Molecular detection of hepatitis E virus in sewage samples","volume":"76","author":"Pourshaban","year":"2010","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"197710","DOI":"10.1016\/j.virusres.2019.197710","article-title":"High prevalence of hepatitis E virus in raw sewage in Southern Italy","volume":"272","author":"Melegari","year":"2019","journal-title":"Virus Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/s12560-012-9097-6","article-title":"Shellfish-borne viral outbreaks: A systematic review","volume":"5","author":"Bellou","year":"2013","journal-title":"Food Environ. Virol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1007\/s12560-016-9228-6","article-title":"Prevalence and Molecular Genotyping of Noroviruses in Market Oysters, Mussels, and Cockles in Bangkok, Thailand","volume":"8","author":"Kittigul","year":"2016","journal-title":"Food Environ. Virol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.fm.2016.05.009","article-title":"Detection and molecular characterization of norovirus from oysters implicated in outbreaks in the US","volume":"59","author":"Woods","year":"2016","journal-title":"Food Microbiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"7350","DOI":"10.1128\/AEM.01177-09","article-title":"Risk assessment in shellfish-borne outbreaks of hepatitis A","volume":"75","author":"Pinto","year":"2009","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"5812","DOI":"10.1128\/AEM.01046-12","article-title":"Two-year systematic study to assess norovirus contamination in oysters from commercial harvesting areas in the United Kingdom","volume":"78","author":"Lowther","year":"2012","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"838","DOI":"10.1086\/511874","article-title":"Use of molecular epidemiology to confirm a multistate outbreak of hepatitis A caused by consumption of oysters","volume":"44","author":"Bialek","year":"2007","journal-title":"Clin. Infect. Dis."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2106081","DOI":"10.1080\/21645515.2022.2106081","article-title":"A community-wide epidemic of hepatitis A virus genotype IA associated with consumption of shellfish in Yantai, eastern China, January to March 2020","volume":"18","author":"Yan","year":"2022","journal-title":"Hum. Vaccin Immunother."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2085","DOI":"10.3201\/eid1812.120924","article-title":"Hepatitis E virus genotype 3 in shellfish, United Kingdom","volume":"18","author":"Crossan","year":"2012","journal-title":"Emerg. Infect. Dis."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1007\/s12560-018-9337-5","article-title":"First Report of the Presence of Hepatitis E Virus in Scottish-Harvested Shellfish Purchased at Retail Level","volume":"10","author":"Crossan","year":"2018","journal-title":"Food Environ. Virol."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Rivadulla, E., Varela, M.F., Mesquita, J.R., Nascimento, M.S.J., and Romalde, J.L. (2019). Detection of Hepatitis E Virus in Shellfish Harvesting Areas from Galicia (Northwestern Spain). Viruses, 11.","DOI":"10.3390\/v11070618"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2026","DOI":"10.3390\/ijerph120202026","article-title":"Surveillance of hepatitis E virus contamination in shellfish in China","volume":"12","author":"Gao","year":"2015","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1738","DOI":"10.3201\/eid1511.091094","article-title":"Hepatitis E outbreak on cruise ship","volume":"15","author":"Said","year":"2009","journal-title":"Emerg. Infect. Dis."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1086","DOI":"10.1111\/j.1365-2036.2008.03679.x","article-title":"A national survey of acute hepatitis E in France","volume":"27","author":"Renou","year":"2008","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1016\/j.jinf.2019.02.001","article-title":"A case-control study into risk factors for acute hepatitis E in the Netherlands, 2015\u20132017","volume":"78","author":"Tulen","year":"2019","journal-title":"J. Infect."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"3883","DOI":"10.1128\/JCM.42.8.3883-3885.2004","article-title":"Infection of a Japanese patient by genotype 4 hepatitis E virus while traveling in Vietnam","volume":"42","author":"Koizumi","year":"2004","journal-title":"J. Clin. Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"27","DOI":"10.2807\/1560-7917.ES2015.20.19.21127","article-title":"Seroprevalence in blood donors reveals widespread, multi-source exposure to hepatitis E virus, southern France, October 2011","volume":"20","author":"Mansuy","year":"2015","journal-title":"Eurosurveillance"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1002\/hep.28436","article-title":"A nationwide survey of hepatitis E viral infection in French blood donors","volume":"63","author":"Mansuy","year":"2016","journal-title":"Hepatology"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1620\/tjem.235.275","article-title":"Clinical Features of Hepatitis E Virus Infection in Ibaraki, Japan: Autochthonous Hepatitis E and Acute-on-Chronic Liver Failure","volume":"235","author":"Inagaki","year":"2015","journal-title":"Tohoku J. Exp. Med."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1931","DOI":"10.1099\/0022-1317-83-8-1931","article-title":"Identification of two distinct genotypes of hepatitis E virus in a Japanese patient with acute hepatitis who had not travelled abroad","volume":"83","author":"Takahashi","year":"2002","journal-title":"J. Gen. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/BF01720406","article-title":"Acute hepatitis E in Catania (eastern Sicily) 1980\u20131994. The role of hepatitis E virus","volume":"25","author":"Cacopardo","year":"1997","journal-title":"Infection"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"802","DOI":"10.1093\/infdis\/jiz207","article-title":"Epidemiology of Hepatitis E in England and Wales: A 10-Year Retrospective Surveillance Study, 2008\u20132017","volume":"220","author":"Oeser","year":"2019","journal-title":"J. Infect. Dis."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1097\/00004836-200111000-00017","article-title":"Acute hepatitis E virus infection presenting as a prolonged cholestatic jaundice","volume":"33","author":"Mechnik","year":"2001","journal-title":"J. Clin. Gastroenterol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1086\/444396","article-title":"Non-travel-associated hepatitis E in England and Wales: Demographic, clinical, and molecular epidemiological characteristics","volume":"192","author":"Ijaz","year":"2005","journal-title":"J. Infect. Dis."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Desdouits, M., Reynaud, Y., Philippe, C., and Guyader, F.S.L. (2023). A Comprehensive Review for the Surveillance of Human Pathogenic Microorganisms in Shellfish. Microorganisms, 11.","DOI":"10.3390\/microorganisms11092218"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1007\/s12560-010-9045-2","article-title":"Processing Strategies to Inactivate Enteric Viruses in Shellfish","volume":"2","author":"Richards","year":"2010","journal-title":"Food Environ. Virol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/S0168-1605(00)00248-8","article-title":"Viruses and bivalve shellfish","volume":"59","author":"Lees","year":"2000","journal-title":"Int. J. Food Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2140","DOI":"10.4315\/0362-028X-70.9.2140","article-title":"Norovirus recognizes histo-blood group antigens on gastrointestinal cells of clams, mussels, and oysters: A possible mechanism of bioaccumulation","volume":"70","author":"Tian","year":"2007","journal-title":"J. Food Prot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"3189","DOI":"10.1128\/AEM.03010-10","article-title":"Strain-dependent norovirus bioaccumulation in oysters","volume":"77","author":"Maalouf","year":"2011","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1783","DOI":"10.4315\/0362-028X.JFP-18-074","article-title":"Presence and Distribution of Histo-Blood Group Antigens in Pacific Oysters and the Effects of Exposure to Noroviruses GI.3 and GII.4 on Their Expression","volume":"81","author":"Su","year":"2018","journal-title":"J. Food Prot."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1128\/AEM.66.4.1375-1378.2000","article-title":"Selective accumulation may account for shellfish-associated viral illness","volume":"66","author":"Burkhardt","year":"2000","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1111\/j.1472-765X.2008.02424.x","article-title":"Occurrence of enteric viruses in shellfish and relation to climatic-environmental factors","volume":"47","author":"Suffredini","year":"2008","journal-title":"Lett. Appl. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"692","DOI":"10.1111\/1541-4337.12271","article-title":"Depuration and Relaying: A Review on Potential Removal of Norovirus from Oysters","volume":"16","author":"McLeod","year":"2017","journal-title":"Compr. Rev. Food Sci. Food Saf."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"McMenemy, P., Kleczkowski, A., Lees, D.N., Lowther, J., and Taylor, N. (2018). A model for estimating pathogen variability in shellfish and predicting minimum depuration times. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0193865"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1198","DOI":"10.3201\/eid1908.130465","article-title":"Norovirus disease in the United States","volume":"19","author":"Hall","year":"2013","journal-title":"Emerg. Infect. Dis."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"4148","DOI":"10.1128\/JCM.40.11.4148-4155.2002","article-title":"Molecular characterization of hepatitis a virus isolates from a transcontinental shellfish-borne outbreak","volume":"40","author":"Sanchez","year":"2002","journal-title":"J. Clin. Microbiol."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Polo, D., Schaeffer, J., Teunis, P., Buchet, V., and Le Guyader, F.S. (2018). Infectivity and RNA Persistence of a Norovirus Surrogate, the Tulane Virus, in Oysters. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.00716"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1007\/s12560-023-09577-w","article-title":"Persistence of Infectious Human Norovirus in Estuarine Water","volume":"16","author":"Rexin","year":"2024","journal-title":"Food Environ. Virol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1038\/nature12029","article-title":"A pathogenic picornavirus acquires an envelope by hijacking cellular membranes","volume":"496","author":"Feng","year":"2013","journal-title":"Nature"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.1111\/jam.14727","article-title":"Hepatitis A infections from food","volume":"129","author":"Randazzo","year":"2020","journal-title":"J. Appl. Microbiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1126\/science.aaf5211","article-title":"Replication of human noroviruses in stem cell-derived human enteroids","volume":"353","author":"Ettayebi","year":"2016","journal-title":"Science"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"6884","DOI":"10.1128\/AEM.05806-11","article-title":"Norovirus infectivity in humans and persistence in water","volume":"77","author":"Seitz","year":"2011","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1097\/MOG.0b013e32835ff238","article-title":"Hepatitis E virus infection","volume":"29","author":"Kamar","year":"2013","journal-title":"Curr. Opin. Gastroenterol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Salvador, D., Neto, C., Benoliel, M.J., and Caeiro, M.F. (2020). Assessment of the Presence of Hepatitis E virus in Surface Water and Drinking Water in Portugal. Microorganisms, 8.","DOI":"10.3390\/microorganisms8050761"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"10042","DOI":"10.1038\/s41598-019-46475-3","article-title":"Population density and water balance influence the global occurrence of hepatitis E epidemics","volume":"9","author":"Carratala","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Ianiro, G., Monini, M., De Sabato, L., Chelli, E., Cerini, N., Ostanello, F., and Di Bartolo, I. (2022). Dynamic of Hepatitis E Virus (HEV) Shedding in Pigs. Animals, 12.","DOI":"10.3390\/ani12091063"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1186\/s40813-021-00189-z","article-title":"Infection dynamics and persistence of hepatitis E virus on pig farms\u2014A review","volume":"7","author":"Meester","year":"2021","journal-title":"Porc. Health Manag."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"4269","DOI":"10.1128\/AEM.00978-14","article-title":"Bioaccumulation efficiency, tissue distribution, and environmental occurrence of hepatitis E virus in bivalve shellfish from France","volume":"80","author":"Grodzki","year":"2014","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Munoz-Chimeno, M., Rodriguez-Paredes, V., Garcia-Lugo, M.A., and Avellon, A. (2022). Hepatitis E genotype 3 genome: A comprehensive analysis of entropy, motif conservation, relevant mutations, and clade-associated polymorphisms. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.1011662"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.fm.2016.03.009","article-title":"Hepatitis E virus genotype 3 in mussels (Mytilus galloprovinciallis), Spain","volume":"58","author":"Mesquita","year":"2016","journal-title":"Food Microbiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1007\/s12560-017-9319-z","article-title":"First Detection of Hepatitis E Virus in Shellfish and in Seawater from Production Areas in Southern Italy","volume":"10","author":"Proroga","year":"2018","journal-title":"Food Environ. Virol."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"La Bella, G., Basanisi, M.G., Nobili, G., Terio, V., Suffredini, E., and La Salandra, G. (2021). First Report of Hepatitis E Virus in Shellfish in Southeast Italy. Appl. Sci., 11.","DOI":"10.3390\/app11010043"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1016\/j.jviromet.2012.06.007","article-title":"Presence of hepatitis E RNA in mussels used as bio-monitors of viral marine pollution","volume":"186","author":"Donia","year":"2012","journal-title":"J. Virol. Methods"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Fusco, G., Anastasio, A., Kingsley, D.H., Amoroso, M.G., Pepe, T., Fratamico, P.M., Cioffi, B., Rossi, R., La Rosa, G., and Boccia, F. (2019). Detection of Hepatitis A Virus and Other Enteric Viruses in Shellfish Collected in the Gulf of Naples, Italy. Int. J. Environ. Res. Public Health, 16.","DOI":"10.3390\/ijerph16142588"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1016\/j.ijfoodmicro.2014.06.012","article-title":"Enteric porcine viruses in farmed shellfish in Denmark","volume":"186","author":"Krog","year":"2014","journal-title":"Int. J. Food Microbiol."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Rastar-Tangeten, M., Mader, M., Schmiedel, S., Weidemann, S., Chalissery, S., Kempski, J., Staufenberger, T., Mazaheri, O., Brandner, J.M., and Addo, M.M. (2025). Lack of evidence for HEV infection in Baltic sea mussels (Mytilidae). BMC Infect. Dis., 25.","DOI":"10.1186\/s12879-025-10727-7"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/s12560-012-9087-8","article-title":"Occurrence of human enteric viruses in commercial mussels at retail level in three European countries","volume":"4","author":"Kokkinos","year":"2012","journal-title":"Food Environ. Virol."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Wei, M.L., Wang, J.F., Wang, Y., Liu, L.B., Xu, X.D., and Wang, J.C. (2025). Development and Application of a Multiplex Reverse Transcription-Droplet Digital PCR Assay for Simultaneous Detection of Hepatitis A Virus and Hepatitis E Virus in Bivalve Shellfish. Foods, 14.","DOI":"10.3390\/foods14010002"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1007\/s12560-025-09661-3","article-title":"Simultaneous Detection of HEV, HAstV and SaV in Bivalve Shellfish Using a Novel Real-Time RT-qPCR Method","volume":"17","author":"Wang","year":"2025","journal-title":"Food Environ. Virol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1002\/jmv.21730","article-title":"Analysis of complete genome sequences of swine hepatitis E virus and possible risk factors for transmission of HEV to humans in Korea","volume":"82","author":"Song","year":"2010","journal-title":"J. Med. Virol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"170","DOI":"10.4269\/ajtmh.2007.76.170","article-title":"Detection of hepatitis E virus RNA from the bivalve Yamato-Shijimi (Corbicula japonica) in Japan","volume":"76","author":"Li","year":"2007","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"2363","DOI":"10.1007\/s00705-012-1422-8","article-title":"Detection and molecular characterization of hepatitis E virus in clinical, environmental and putative animal sources","volume":"157","author":"Ishida","year":"2012","journal-title":"Arch. Virol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"608","DOI":"10.1111\/j.1472-765X.2011.03152.x","article-title":"Surveillance of hepatitis A and E viruses contamination in shellfish in Thailand","volume":"53","author":"Namsai","year":"2011","journal-title":"Lett. Appl. Microbiol."},{"key":"ref_86","first-page":"7401","article-title":"Hepatitis E virus detection in shellfish using real-time PCR analysis from Oualidia Lagoon in Morocco","volume":"45","author":"Bazir","year":"2022","journal-title":"Teikyo Med. J."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1007\/s12560-024-09624-0","article-title":"Detection of Rocahepevirus ratti in Bivalve Mollusks from Sao Luis Island, Maranhao, Brazil: A Potential Transmission Route of an Emerging Zoonotic Pathogen?","volume":"17","author":"Figueiredo","year":"2025","journal-title":"Food Environ. Virol."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"EFSA Panel on Biological Hazards (BIOHAZ), Ricci, A., Allende, A., Bolton, D., Chemaly, M., Davies, R., Fernandez Escamez, P.S., Herman, L., Koutsoumanis, K., and Lindqvist, R. (2017). Public health risks associated with hepatitis E virus (HEV) as a food-borne pathogen. EFSA J., 15, e04886.","DOI":"10.2903\/j.efsa.2017.4886"},{"key":"ref_89","first-page":"e02908","article-title":"Assessing the incidence of enteric viruses in bivalve mollusks at Alexandria\u2019s seafood markets","volume":"29","author":"Ahmed","year":"2025","journal-title":"Sci. Afr."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s12560-020-09456-8","article-title":"Occurrence of Human Enteric Viruses in Water Sources and Shellfish: A Focus on Africa","volume":"13","author":"Upfold","year":"2021","journal-title":"Food Environ. Virol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"11786302241299370","DOI":"10.1177\/11786302241299370","article-title":"Zoonotic and Food-Related Hazards Due to Hepatitis A and E in Africa: A Systematic Review and Meta-Analysis","volume":"18","author":"Odoom","year":"2024","journal-title":"Environ. Health Insights"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/s12560-025-09639-1","article-title":"Occurrence and Molecular Characterization of Human Astrovirus and Hepatitis A Virus in Bivalve Mollusks Marketed in Tourist Cities in Rio de Janeiro, Brazil","volume":"17","author":"Burlandy","year":"2025","journal-title":"Food Environ. Virol."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"do Nascimento, L.G., Sarmento, S.K., Leonardo, R., Gutierrez, M.B., Malta, F.C., de Oliveira, J.M., Guerra, C.R., Coutinho, R., Miagostovich, M.P., and Fumian, T.M. (2022). Detection and Molecular Characterization of Enteric Viruses in Bivalve Mollusks Collected in Arraial do Cabo, Rio de Janeiro, Brazil. Viruses, 14.","DOI":"10.3390\/v14112359"},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Viera-Segura, O., Calderon-Flores, A., Batun-Alfaro, J.A., and Fierro, N.A. (2023). Tracing the History of Hepatitis E Virus Infection in Mexico: From the Enigmatic Genotype 2 to the Current Disease Situation. Viruses, 15.","DOI":"10.3390\/v15091911"},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"European Association for the Study of the Liver (2018). EASL Clinical Practice Guidelines on hepatitis E virus infection. J. Hepatol., 68, 1256\u20131271.","DOI":"10.1016\/j.jhep.2018.03.005"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/hepr.14062","article-title":"Recent advances in hepatitis E virus research and the Japanese clinical practice guidelines for hepatitis E virus infection","volume":"54","author":"Kanda","year":"2024","journal-title":"Hepatol. Res."},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Lin, D., Chen, W., Lin, Z., Liu, L., Zhang, M., Yang, H., Liu, Z., and Chen, L. (2025). Viral Transmission in Sea Food Systems: Strategies for Control and Emerging Challenges. Foods, 14.","DOI":"10.3390\/foods14061071"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"5153","DOI":"10.1128\/AEM.00436-12","article-title":"Thermal inactivation of infectious hepatitis E virus in experimentally contaminated food","volume":"78","author":"Barnaud","year":"2012","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.ijfoodmicro.2007.11.068","article-title":"Inactivation of infectious hepatitis E virus present in commercial pig livers sold in local grocery stores in the United States","volume":"123","author":"Feagins","year":"2008","journal-title":"Int. J. Food Microbiol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1163","DOI":"10.1111\/jam.13242","article-title":"Efficacy of domestic cooking inactivation of human hepatitis A virus in experimentally infected manila clams (Ruditapes philippinarum)","volume":"121","author":"Pascoli","year":"2016","journal-title":"J. Appl. Microbiol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1111\/jam.12892","article-title":"Thermal inactivation kinetics of hepatitis A virus in homogenized clam meat (Mercenaria mercenaria)","volume":"119","author":"Bozkurt","year":"2015","journal-title":"J. Appl. Microbiol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1007\/s12560-010-9052-3","article-title":"Heat Inactivation of Hepatitis A Virus in Shellfish Using Steam","volume":"3","author":"Harlow","year":"2011","journal-title":"Food Environ. Virol."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1021\/acs.estlett.2c00553","article-title":"Viable Norovirus Persistence in Water Microcosms","volume":"9","author":"Shaffer","year":"2022","journal-title":"Environ. Sci. Technol. Lett."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1093\/infdis\/jiu385","article-title":"Two Human Challenge Studies Confirm High Infectivity of Norwalk Virus","volume":"211","author":"Kirby","year":"2015","journal-title":"J. Infect. Dis."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1002\/rmv.678","article-title":"Efficient cell culture systems for hepatitis E virus strains in feces and circulating blood","volume":"21","author":"Okamoto","year":"2011","journal-title":"Rev. Med. Virol."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Zhang, J., Liang, Z., Liu, F., Wang, Y., Huang, W., and Nie, J. (2025). Research Progress on Hepatitis E Virus Culture. Pathogens, 14.","DOI":"10.3390\/pathogens14050456"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.antiviral.2019.01.007","article-title":"Cell culture systems for the study of hepatitis E virus","volume":"163","author":"Meister","year":"2019","journal-title":"Antivir. Res."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Fu, R.M., Decker, C.C., and Dao Thi, V.L. (2019). Cell Culture Models for Hepatitis E Virus. Viruses, 11.","DOI":"10.3390\/v11070608"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1099\/jgv.0.000393","article-title":"Proposed reference sequences for hepatitis E virus subtypes","volume":"97","author":"Smith","year":"2016","journal-title":"J. Gen. Virol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.2147\/IDR.S449221","article-title":"Hepatitis E Virus (HEV) Infection Among Immunocompromised Individuals: A Brief Narrative Review","volume":"17","author":"Alexandrova","year":"2024","journal-title":"Infect. 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