{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,9,11]],"date-time":"2025-09-11T18:09:42Z","timestamp":1757614182353,"version":"3.44.0"},"reference-count":75,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,9,5]],"date-time":"2025-09-05T00:00:00Z","timestamp":1757030400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"abstract":"<jats:sec><jats:title>Background<\/jats:title><jats:p>Hepatitis E virus (HEV) particles are released from infected cells in a quasi-enveloped form, typically via the multivesicular body (MVB) pathway, which is mediated by the viral accessory protein pORF3. However, cell-type specific aspects of this release mechanism remain poorly understood.<\/jats:p><\/jats:sec><jats:sec><jats:title>Methods<\/jats:title><jats:p>We analyzed the release and envelopment characteristics of a pORF3-deficient genotype 3c HEV (HEV\u0394ORF3) in comparison to wild-type HEV (HEVwt) in two human cell lines: hepatoma-derived PLC\/PRF\/5 and lung carcinoma-derived A549\/D3 cells.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>While viral release of HEV\u0394ORF3 was strongly impaired in A549\/D3 cells, PLC\/PRF\/5 cells supported efficient viral release despite the absence of pORF3. In PLC\/PRF\/5 cells, HEV particles retained quasi-envelopment and utilized an alternative, Golgi-associated egress pathway in the absence of pORF3. In contrast, A549\/D3 cells did not support this compensatory release route.<\/jats:p><\/jats:sec><jats:sec><jats:title>Conclusion<\/jats:title><jats:p>Our findings highlight a pronounced cell line-dependent variability in HEV release pathways, emphasizing the importance of cellular context in studies of HEV biology and antiviral strategies targeting virus egress.<\/jats:p><\/jats:sec>","DOI":"10.3389\/fcimb.2025.1661270","type":"journal-article","created":{"date-parts":[[2025,9,5]],"date-time":"2025-09-05T05:32:13Z","timestamp":1757050333000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Cell line-dependent release of quasi-enveloped hepatitis E virus reveals alternative Golgi-associated egress in the absence of pORF3"],"prefix":"10.3389","volume":"15","author":[{"given":"Nele","family":"Gremmel","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mirco","family":"Glitscher","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Johannes","family":"Scholz","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ashish","family":"Gadicherla","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Reimar","family":"Johne","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eberhard","family":"Hildt","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Paul","family":"Becher","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2025,9,5]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"6243","DOI":"10.1038\/s41598-019-42737-2","article-title":"New insights into the ORF2 capsid protein, a key player of the hepatitis E virus lifecycle","volume":"9","author":"Ankavay","year":"2019","journal-title":"Sci. Rep."},{"key":"B2","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1007\/s00018-022-04646-y","article-title":"The endocytic recycling compartment serves as a viral factory for hepatitis E virus","volume":"79","author":"Bentaleb","year":"2022","journal-title":"Cell Mol. Life Sci."},{"key":"B3","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1016\/j.jhep.2016.04.013","article-title":"First-in-human application of the novel hepatitis B and hepatitis D virus entry inhibitor myrcludex B","volume":"65","author":"Blank","year":"2016","journal-title":"J. Hepatol."},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.01207-18","article-title":"Vectorial release of hepatitis E virus in polarized human hepatocytes","volume":"93","author":"Capelli","year":"2019","journal-title":"J. Virol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1038\/286531a0","article-title":"Identification of integrated hepatitis B virus DNA and expression of viral RNA in an HBsAg-producing human hepatocellular carcinoma cell line","volume":"286","author":"Chakraborty","year":"1980","journal-title":"Nature"},{"key":"B6","doi-asserted-by":"publisher","first-page":"326","DOI":"10.1007\/s00018-023-04977-4","article-title":"The phosphatidylserine receptor TIM1 promotes infection of enveloped hepatitis E virus","volume":"80","author":"Corneillie","year":"2023","journal-title":"Cell Mol. Life Sci."},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.3390\/v15122412","article-title":"Virus-host protein interaction network of the hepatitis E virus ORF2\u20134 by mammalian two-hybrid assays","volume":"15","author":"Corneillie","year":"2023","journal-title":"Viruses"},{"key":"B8","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1038\/s41579-023-00889-z","article-title":"Cell entry and release of quasi-enveloped human hepatitis viruses","volume":"21","author":"Das","year":"2023","journal-title":"Nat. Rev. Microbiol."},{"key":"B9","doi-asserted-by":"publisher","first-page":"1147","DOI":"10.1073\/pnas.1614955114","article-title":"Hepatitis E virus ORF3 is a functional ion channel required for release of infectious particles","volume":"114","author":"Ding","year":"2017","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B10","doi-asserted-by":"publisher","first-page":"9059","DOI":"10.1128\/JVI.00593-10","article-title":"Release of genotype 1 hepatitis E virus from cultured hepatoma and polarized intestinal cells depends on open reading frame 3 protein and requires an intact PXXP motif","volume":"84","author":"Emerson","year":"2010","journal-title":"J. Virol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"10457","DOI":"10.1128\/JVI.00892-06","article-title":"ORF3 protein of hepatitis E virus is not required for replication, virion assembly, or infection of hepatoma cells in vitro","volume":"80","author":"Emerson","year":"2006","journal-title":"J. Virol."},{"key":"B12","doi-asserted-by":"publisher","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":"B13","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1007\/s00018-024-05367-0","article-title":"The AP-1 adaptor complex is essential for intracellular trafficking of the ORF2 capsid protein and assembly of Hepatitis E virus","volume":"81","author":"Ferri\u00e9","year":"2024","journal-title":"Cell Mol. Life Sci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"16","DOI":"10.3350\/cmh.2019.0082","article-title":"Extrahepatic manifestations of hepatitis E virus: An overview","volume":"26","author":"Fousekis","year":"2020","journal-title":"Clin. Mol. Hepatol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"G432","DOI":"10.1152\/ajpgi.00109.2020","article-title":"Acetaldehyde suppresses HBV-MHC class I complex presentation on hepatocytes via induction of ER stress and Golgi fragmentation","volume":"319","author":"Ganesan","year":"2020","journal-title":"Am. J. Physiology-Gastrointestinal Liver Physiol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.micinf.2018.06.002","article-title":"Organelle dynamics and viral infections: at cross roads","volume":"21","author":"Glingston","year":"2019","journal-title":"Microbes Infect."},{"key":"B17","doi-asserted-by":"publisher","first-page":"e13379","DOI":"10.1111\/cmi.13379","article-title":"Hepatitis E virus egress and beyond - the manifold roles of the viral ORF3 protein","volume":"23","author":"Glitscher","year":"2021","journal-title":"Cell Microbiol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"e01564","DOI":"10.1128\/JVI.01564-20","article-title":"Identification of the interferon-inducible GTPase GBP1 as major restriction factor for the Hepatitis E virus","volume":"95","author":"Glitscher","year":"2021","journal-title":"J. Virol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/j.jcmgh.2021.02.002","article-title":"Targeting cholesterol metabolism as efficient antiviral strategy against the hepatitis E virus","volume":"12","author":"Glitscher","year":"2021","journal-title":"Cell Mol. Gastroenterol. Hepatol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1016\/j.jcmgh.2024.01.001","article-title":"The protease domain in HEV pORF1 mediates the replicase\u2019s localization to multivesicular bodies and its exosomal release","volume":"17","author":"Glitscher","year":"2024","journal-title":"Cell Mol. Gastroenterol. Hepatol."},{"key":"B21","doi-asserted-by":"publisher","first-page":"e1007471","DOI":"10.1371\/journal.ppat.1007471","article-title":"Palmitoylation mediates membrane association of hepatitis E virus ORF3 protein and is required for infectious particle secretion","volume":"14","author":"Gouttenoire","year":"2018","journal-title":"PLoS Pathog."},{"key":"B22","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1128\/JVI.01219-07","article-title":"Mutations within potential glycosylation sites in the capsid protein of hepatitis E virus prevent the formation of infectious virus particles","volume":"82","author":"Graff","year":"2008","journal-title":"J. Virol."},{"key":"B23","doi-asserted-by":"publisher","first-page":"e2617","DOI":"10.1111\/tbed.14608","article-title":"Isolation of 15 hepatitis E virus strains lacking ORF1 rearrangements from wild boar and pig organ samples and efficient replication in cell culture","volume":"69","author":"Gremmel","year":"2022","journal-title":"Transbound Emerg. Dis."},{"key":"B24","doi-asserted-by":"publisher","first-page":"e00373","DOI":"10.1128\/jcm.00373-23","article-title":"Hepatitis E virus neutralization by porcine serum antibodies","volume":"61","author":"Gremmel","year":"2023","journal-title":"J. Clin. Microbiol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"905","DOI":"10.1038\/s41564-024-01637-6","article-title":"Phosphatidylserine-exposing extracellular vesicles in body fluids are an innate defence against apoptotic mimicry viral pathogens","volume":"9","author":"Gro\u00df","year":"2024","journal-title":"Nat. Microbiol."},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1616683114","article-title":"Hepatitis C virus triggers Golgi fragmentation and autophagy through the immunity-related GTPase M","volume":"114","author":"Hansen","year":"2017","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1038\/srep27597","article-title":"The ORF3 protein of genotype 1 hepatitis E virus suppresses TLR3-induced NF-\u03baB signaling via TRADD and RIP1","volume":"6","author":"He","year":"2016","journal-title":"Sci. Rep."},{"key":"B28","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/j.jhep.2020.12.030","article-title":"Hepatitis E virus persists in the ejaculate of chronically infected men","volume":"75","author":"Horvatits","year":"2021","journal-title":"J. Hepatol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1016\/S0140-6736(23)02234-1","article-title":"Long-term efficacy of a recombinant hepatitis E vaccine in adults: 10-year results from a randomised, double-blind, placebo-controlled, phase 3 trial","volume":"403","author":"Huang","year":"2024","journal-title":"Lancet"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.609516","article-title":"Differentiation of trafficking pathways at golgi entry core compartments and post-golgi subdomains","volume":"11","author":"Ito","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B31","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1128\/JVI.70.1.207-216.1996","article-title":"Expression in animal cells and characterization of the hepatitis E virus structural proteins","volume":"70","author":"Jameel","year":"1996","journal-title":"J. Virol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"447","DOI":"10.1111\/jvh.12157","article-title":"An ORF1-rearranged hepatitis E virus derived from a chronically infected patient efficiently replicates in cell culture","volume":"21","author":"Johne","year":"2014","journal-title":"J. Viral Hepat"},{"key":"B33","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/j.jviromet.2005.07.004","article-title":"A broadly reactive one-step real-time RT-PCR assay for rapid and sensitive detection of hepatitis E virus","volume":"131","author":"Jothikumar","year":"2006","journal-title":"J. Virol. Methods"},{"key":"B34","doi-asserted-by":"publisher","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":"B35","doi-asserted-by":"publisher","first-page":"5637","DOI":"10.1128\/JVI.06711-11","article-title":"The PSAP motif within the ORF3 protein of an avian strain of the hepatitis E virus is not critical for viral infectivity in vivo but plays a role in virus release","volume":"86","author":"Kenney","year":"2012","journal-title":"J. Virol."},{"key":"B36","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1016\/j.virol.2015.09.012","article-title":"Replacement of the hepatitis E virus ORF3 protein PxxP motif with heterologous late domain motifs affects virus release via interaction with TSG101","volume":"486","author":"Kenney","year":"2015","journal-title":"Virology"},{"key":"B37","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1053\/j.gastro.2015.10.048","article-title":"Chronic infection with camelid hepatitis E virus in a liver transplant recipient who regularly consumes camel meat and milk","volume":"150","author":"Lee","year":"2016","journal-title":"Gastroenterology"},{"key":"B38","doi-asserted-by":"publisher","first-page":"8585","DOI":"10.1038\/s41598-018-26975-4","article-title":"HEV ORF3 downregulates TLR7 to inhibit the generation of type I interferon via impairment of multiple signaling pathways","volume":"8","author":"Lei","year":"2018","journal-title":"Sci. Rep."},{"key":"B39","doi-asserted-by":"publisher","first-page":"1516","DOI":"10.1111\/liv.14468","article-title":"The global epidemiology of hepatitis E virus infection: A systematic review and meta-analysis","volume":"40","author":"Li","year":"2020","journal-title":"Liver Int."},{"key":"B40","doi-asserted-by":"publisher","first-page":"50","DOI":"10.4161\/cib.18208","article-title":"Vesicle formation within endosomes: An ESCRT marks the spot","volume":"5","author":"Mayers","year":"2012","journal-title":"Commun. Integr. Biol."},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2022.828636","article-title":"Processing and subcellular localization of the hepatitis E virus replicase: identification of candidate viral factories","volume":"13","author":"Metzger","year":"2022","journal-title":"Front. Microbiol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1053\/j.gastro.2017.09.020","article-title":"Hepatitis E virus lifecycle and identification of 3 forms of the ORF2 capsid protein","volume":"154","author":"Montpellier","year":"2018","journal-title":"Gastroenterology"},{"key":"B43","doi-asserted-by":"publisher","first-page":"2166","DOI":"10.1099\/vir.0.066910-0","article-title":"Hepatitis E virus egress depends on the exosomal pathway, with secretory exosomes derived from multivesicular bodies","volume":"95","author":"Nagashima","year":"2014","journal-title":"J. Gen. Virol."},{"key":"B44","doi-asserted-by":"publisher","first-page":"979","DOI":"10.1007\/s00705-013-1912-3","article-title":"The membrane on the surface of hepatitis E virus particles is derived from the intracellular membrane and contains trans-Golgi network protein 2","volume":"159","author":"Nagashima","year":"2014","journal-title":"Arch. Virol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"2838","DOI":"10.1099\/vir.0.035378-0","article-title":"Tumour susceptibility gene 101 and the vacuolar protein sorting pathway are required for the release of hepatitis E virions","volume":"92","author":"Nagashima","year":"2011","journal-title":"J. Gen. Virol."},{"key":"B46","doi-asserted-by":"publisher","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":"B47","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1186\/s13567-017-0483-9","article-title":"Recent knowledge on hepatitis E virus in Suidae reservoirs and transmission routes to human","volume":"48","author":"Pavio","year":"2017","journal-title":"Vet. Res."},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1016\/j.virusres.2020.197868","article-title":"Multivesicular body sorting and the exosomal pathway are required for the release of rat hepatitis E virus from infected cells","volume":"278","author":"Primadharsini","year":"2020","journal-title":"Virus Res."},{"key":"B49","doi-asserted-by":"publisher","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":"B50","doi-asserted-by":"publisher","first-page":"e0132503","DOI":"10.1371\/journal.pone.0132503","article-title":"Hepatitis E virus produced from cell culture has a lipid envelope","volume":"10","author":"Qi","year":"2015","journal-title":"PLoS One"},{"key":"B51","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1016\/j.ceb.2019.10.003","article-title":"The Golgi apparatus and cell polarity: Roles of the cytoskeleton, the Golgi matrix, and Golgi membranes","volume":"62","author":"Ravichandran","year":"2020","journal-title":"Curr. Opin. Cell Biol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1016\/0042-6822(92)90716-3","article-title":"Indian hepatitis E virus shows a major deletion in the small open reading frame","volume":"189","author":"Ray","year":"1992","journal-title":"Virology"},{"key":"B53","doi-asserted-by":"publisher","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":"B54","doi-asserted-by":"publisher","first-page":"208","DOI":"10.1016\/j.chom.2018.07.006","article-title":"Vesicle-cloaked virus clusters are optimal units for inter-organismal viral transmission","volume":"24","author":"Santiana","year":"2018","journal-title":"Cell Host Microbe"},{"key":"B55","doi-asserted-by":"publisher","first-page":"e00585","DOI":"10.1128\/JVI.00585-21","article-title":"The viral ORF3 protein is required for hepatitis E virus apical release and efficient growth in polarized hepatocytes and humanized mice","volume":"95","author":"Sari","year":"2021","journal-title":"J. Virol."},{"key":"B56","doi-asserted-by":"publisher","first-page":"267","DOI":"10.3390\/v8100267","article-title":"Enhanced replication of hepatitis E virus strain 47832c in an A549-derived subclonal cell line","volume":"8","author":"Schemmerer","year":"2016","journal-title":"Viruses"},{"key":"B57","doi-asserted-by":"publisher","first-page":"483","DOI":"10.3390\/v11060483","article-title":"Isolation of subtype 3c, 3e and 3f-like hepatitis E virus strains stably replicating to high viral loads in an optimized cell culture system","volume":"11","author":"Schemmerer","year":"2019","journal-title":"Viruses"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms9112302","article-title":"Cell culture isolation and whole genome characterization of hepatitis E virus strains from wild boars in Germany","volume":"9","author":"Schilling-Loeffler","year":"2021","journal-title":"Microorganisms"},{"key":"B59","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"B60","doi-asserted-by":"publisher","first-page":"R116","DOI":"10.1016\/j.cub.2012.01.028","article-title":"The ESCRT machinery","volume":"22","author":"Schmidt","year":"2012","journal-title":"Curr. Biol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"157","DOI":"10.3390\/pathogens9030157","article-title":"Establishment of a plasmid-based reverse genetics system for the cell culture-adapted hepatitis E virus genotype 3c strain 47832c","volume":"9","author":"Scholz","year":"2020","journal-title":"Pathogens"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1016\/j.antiviral.2024.105940","article-title":"Zika virus replication is impaired by a selective agonist of the TRPML2 ion channel","volume":"228","author":"Schwickert","year":"2024","journal-title":"Antiviral Res."},{"key":"B63","doi-asserted-by":"publisher","first-page":"e00135","DOI":"10.1128\/mSystems.00135-17","article-title":"Host-virus protein interaction network reveals the involvement of multiple host processes in the life cycle of hepatitis E virus","volume":"3","author":"Subramani","year":"2018","journal-title":"mSystems"},{"key":"B64","doi-asserted-by":"publisher","first-page":"320","DOI":"10.1128\/jvi.78.1.320-328.2004","article-title":"The ORF2 protein of hepatitis E virus binds the 5\u2019 region of viral RNA","volume":"78","author":"Surjit","year":"2004","journal-title":"J. Virol."},{"key":"B65","doi-asserted-by":"publisher","first-page":"8135","DOI":"10.1074\/jbc.M509568200","article-title":"Enhanced \u03b11 microglobulin secretion from hepatitis E virus ORF3-expressing human hepatoma cells is mediated by the tumor susceptibility gene 101*","volume":"281","author":"Surjit","year":"2006","journal-title":"J. Biol. Chem."},{"key":"B66","doi-asserted-by":"publisher","first-page":"e00459","DOI":"10.1128\/JVI.00459-19","article-title":"Recombinant hepatitis E viruses harboring tags in the ORF1 protein","volume":"93","author":"Szkolnicka","year":"2019","journal-title":"J. Virol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"1703","DOI":"10.1007\/s00705-008-0179-6","article-title":"Monoclonal antibodies raised against the ORF3 protein of hepatitis E virus (HEV) can capture HEV particles in culture supernatant and serum but not those in feces","volume":"153","author":"Takahashi","year":"2008","journal-title":"Arch. Virol."},{"key":"B68","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1021\/cb800193n","article-title":"Inhibition of HIV budding by a genetically selected cyclic peptide targeting the Gag-TSG101 interaction","volume":"3","author":"Tavassoli","year":"2008","journal-title":"ACS Chem. Biol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"22759","DOI":"10.1074\/jbc.M200185200","article-title":"The phosphorylated form of the ORF3 protein of hepatitis E virus interacts with its non-glycosylated form of the major capsid protein, ORF2*","volume":"277","author":"Tyagi","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B70","doi-asserted-by":"publisher","first-page":"e0058024","DOI":"10.1128\/jvi.00580-24","article-title":"Antagonism of epidermal growth factor receptor signaling favors hepatitis E virus life cycle","volume":"98","author":"Woytinek","year":"2024","journal-title":"J. Virol."},{"key":"B71","doi-asserted-by":"publisher","first-page":"33175","DOI":"10.1074\/jbc.M110.106336","article-title":"Structure of hepatitis E virion-sized particle reveals an RNA-dependent viral assembly pathway","volume":"285","author":"Xing","year":"2010","journal-title":"J. Biol. Chem."},{"key":"B72","doi-asserted-by":"publisher","first-page":"1880","DOI":"10.1099\/vir.0.010561-0","article-title":"ORF3 protein of hepatitis E virus is essential for virion release from infected cells","volume":"90","author":"Yamada","year":"2009","journal-title":"J. Gen. Virol."},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.3390\/vetsci9120676","article-title":"Application of ORF3 subunit vaccine for avian hepatitis E virus","volume":"9","author":"Yan","year":"2022","journal-title":"Vet. Sci."},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.3389\/fmolb.2021.703640","article-title":"The key role of exosomes on the pre-metastatic niche formation in tumors","volume":"8","author":"Yang","year":"2021","journal-title":"Front. Mol. Biosci."},{"key":"B75","doi-asserted-by":"publisher","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."}],"container-title":["Frontiers in Cellular and Infection Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2025.1661270\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,5]],"date-time":"2025-09-05T05:32:13Z","timestamp":1757050333000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2025.1661270\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,9,5]]},"references-count":75,"alternative-id":["10.3389\/fcimb.2025.1661270"],"URL":"https:\/\/doi.org\/10.3389\/fcimb.2025.1661270","relation":{},"ISSN":["2235-2988"],"issn-type":[{"type":"electronic","value":"2235-2988"}],"subject":[],"published":{"date-parts":[[2025,9,5]]},"article-number":"1661270"}}