{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T21:30:01Z","timestamp":1790803801221,"version":"4.1.0"},"reference-count":47,"publisher":"Elsevier BV","issue":"11","license":[{"start":{"date-parts":[[2019,11,1]],"date-time":"2019-11-01T00:00:00Z","timestamp":1572566400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2019,11,1]],"date-time":"2019-11-01T00:00:00Z","timestamp":1572566400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2019,11,16]],"date-time":"2019-11-16T00:00:00Z","timestamp":1573862400000},"content-version":"vor","delay-in-days":15,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100009619","name":"Japan Agency for Medical Research and Development","doi-asserted-by":"publisher","award":["JP19fk0108035"],"award-info":[{"award-number":["JP19fk0108035"]}],"id":[{"id":"10.13039\/100009619","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Heliyon"],"published-print":{"date-parts":[[2019,11]]},"DOI":"10.1016\/j.heliyon.2019.e02882","type":"journal-article","created":{"date-parts":[[2019,11,23]],"date-time":"2019-11-23T02:20:15Z","timestamp":1574475615000},"page":"e02882","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":31,"title":["E and prM proteins of genotype V Japanese encephalitis virus are required for its increased virulence in mice"],"prefix":"10.1016","volume":"5","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1981-9062","authenticated-orcid":false,"given":"Shigeru","family":"Tajima","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken-ichi","family":"Shibasaki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Satoshi","family":"Taniguchi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0826-3513","authenticated-orcid":false,"given":"Eri","family":"Nakayama","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takahiro","family":"Maeki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chang-Kweng","family":"Lim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5458-7298","authenticated-orcid":false,"given":"Masayuki","family":"Saijo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.heliyon.2019.e02882_bib1","doi-asserted-by":"crossref","first-page":"e70","DOI":"10.3346\/jkms.2018.33.e70","article-title":"Changes of epidemiological characteristics of Japanese encephalitis viral infection and birds as a potential viral transmitter in Korea","volume":"33","author":"Bae","year":"2018","journal-title":"J.\u00a0Korean Med. Sci."},{"key":"10.1016\/j.heliyon.2019.e02882_bib2","doi-asserted-by":"crossref","first-page":"766","DOI":"10.2471\/BLT.10.085233","article-title":"Estimated global incidence of Japanese encephalitis: a systematic review","volume":"89","author":"Campbell","year":"2011","journal-title":"Bull. World Health Organ."},{"key":"10.1016\/j.heliyon.2019.e02882_bib3","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0004686","article-title":"Low protective efficacy of the current Japanese encephalitis vaccine against the emerging genotype 5 Japanese encephalitis virus","volume":"10","author":"Cao","year":"2016","journal-title":"PLoS Neglected Trop. Dis."},{"key":"10.1016\/j.heliyon.2019.e02882_bib4","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.virol.2007.04.016","article-title":"JE Nakayama\/JE SA14-14-2 virus structural region intertypic viruses: biological properties in the mouse model of neuroinvasive disease","volume":"366","author":"Chambers","year":"2007","journal-title":"Virology"},{"key":"10.1016\/j.heliyon.2019.e02882_bib5","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1002\/jmv.20406","article-title":"Fitness of Japanese encephalitis virus to Neuro-2a cells is determined by interactions of the viral envelope protein with highly sulfated glycosaminoglycans on the cell surface","volume":"76","author":"Chiou","year":"2005","journal-title":"J.\u00a0Med. Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib6","doi-asserted-by":"crossref","first-page":"5862","DOI":"10.1128\/JVI.00358-15","article-title":"A Japanese Encephalitis virus genotype 5 molecular clone is highly neuropathogenic in a mouse model: implication of the structural proteins region in virulence","volume":"89","author":"de Wispelaere","year":"2015","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib7","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3201\/eid1501.080311","article-title":"Past, present, and future of Japanese encephalitis","volume":"15","author":"Erlanger","year":"2009","journal-title":"Emerg. Infect. Dis."},{"key":"10.1016\/j.heliyon.2019.e02882_bib8","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1083\/jcb.200806081","article-title":"Identification of specific histidines as pH sensors in flavivirus membrane fusion","volume":"183","author":"Fritz","year":"2008","journal-title":"J.\u00a0Cell Biol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib9","doi-asserted-by":"crossref","first-page":"e2459","DOI":"10.1371\/journal.pntd.0002459","article-title":"Southernmost Asia is the source of Japanese encephalitis virus (genotype 1) diversity from which the viruses disperse and evolve throughout Asia","volume":"7","author":"Gao","year":"2013","journal-title":"PLoS Neglected Trop. Dis."},{"key":"10.1016\/j.heliyon.2019.e02882_bib10","doi-asserted-by":"crossref","first-page":"6328","DOI":"10.1128\/JVI.00219-15","article-title":"Genetic determinants of Japanese encephalitis virus vaccine strain SA14-14-2 that govern attenuation of virulence in mice","volume":"89","author":"Gromowski","year":"2015","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib11","doi-asserted-by":"crossref","first-page":"16160","DOI":"10.1038\/s41598-017-16475-2","article-title":"Characterization of large and small-plaque variants in the Zika virus clinical isolate ZIKV\/Hu\/S36\/Chiba\/2016","volume":"7","author":"Kato","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.heliyon.2019.e02882_bib12","article-title":"Detection of Japanese encephalitis virus genotype V in Culex orientalis and Culex pipiens (Diptera: Culicidae) in Korea","volume":"10","author":"Kim","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.heliyon.2019.e02882_bib13","doi-asserted-by":"crossref","first-page":"7846","DOI":"10.1128\/JVI.00789-08","article-title":"A\u00a0single N-linked glycosylation site in the Japanese encephalitis virus prM protein is critical for cell type-specific prM protein biogenesis, virus particle release, and pathogenicity in mice","volume":"82","author":"Kim","year":"2008","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib14","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1006\/viro.1999.9859","article-title":"Prediction of three-dimensional structure and mapping of conformational epitopes of envelope glycoprotein of Japanese encephalitis virus","volume":"261","author":"Kolaskar","year":"1999","journal-title":"Virology"},{"key":"10.1016\/j.heliyon.2019.e02882_bib15","doi-asserted-by":"crossref","first-page":"1672","DOI":"10.1128\/jvi.67.3.1672-1675.1993","article-title":"Proper maturation of the Japanese encephalitis virus envelope glycoprotein requires cosynthesis with the premembrane protein","volume":"67","author":"Konishi","year":"1993","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib16","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0001231","article-title":"Genotype v Japanese encephalitis virus is emerging","volume":"5","author":"Li","year":"2011","journal-title":"PLoS Neglected Trop. Dis."},{"key":"10.1016\/j.heliyon.2019.e02882_bib17","doi-asserted-by":"crossref","first-page":"2746","DOI":"10.1016\/j.vaccine.2009.03.007","article-title":"A\u00a0Japanese encephalitis virus vaccine candidate strain is attenuated by decreasing its interferon antagonistic ability","volume":"27","author":"Liang","year":"2009","journal-title":"Vaccine"},{"key":"10.1016\/j.heliyon.2019.e02882_bib18","doi-asserted-by":"crossref","first-page":"5480","DOI":"10.1128\/JVI.76.11.5480-5491.2002","article-title":"Folding and dimerization of tick-borne encephalitis virus envelope proteins prM and E in the endoplasmic reticulum","volume":"76","author":"Lorenz","year":"2002","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib19","doi-asserted-by":"crossref","first-page":"2337","DOI":"10.1128\/JVI.06072-11","article-title":"Crystal structure of the Japanese encephalitis virus envelope protein","volume":"86","author":"Luca","year":"2012","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib20","doi-asserted-by":"crossref","first-page":"1932","DOI":"10.1128\/JVI.76.4.1932-1943.2002","article-title":"Single mutation in the flavivirus envelope protein hinge region increases neurovirulence for mice and monkeys but decreases viscerotropism for monkeys: relevance to development and safety testing of live, attenuated vaccines","volume":"76","author":"Monath","year":"2002","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib21","doi-asserted-by":"crossref","first-page":"3448","DOI":"10.1128\/JVI.79.6.3448-3458.2005","article-title":"Nuclear localization of Japanese encephalitis virus core protein enhances viral replication","volume":"79","author":"Mori","year":"2005","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib22","doi-asserted-by":"crossref","first-page":"12631","DOI":"10.1128\/JVI.01072-09","article-title":"Protonation of individual histidine residues is not required for the pH-dependent entry of west nile virus: evaluation of the \"histidine switch\" hypothesis","volume":"83","author":"Nelson","year":"2009","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib23","doi-asserted-by":"crossref","first-page":"2762","DOI":"10.1099\/vir.0.82941-0","article-title":"Molecular epidemiological analyses of Japanese encephalitis virus isolates from swine in Japan from 2002 to 2004","volume":"88","author":"Nerome","year":"2007","journal-title":"J.\u00a0Gen. Virol."},{"issue":"Pt 7","key":"10.1016\/j.heliyon.2019.e02882_bib24","doi-asserted-by":"crossref","first-page":"1449","DOI":"10.1099\/0022-1317-77-7-1449","article-title":"Molecular differences between wild-type Japanese encephalitis virus strains of high and low mouse neuroinvasiveness","volume":"77","author":"Ni","year":"1996","journal-title":"J.\u00a0Gen. Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib25","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1006\/viro.1997.8956","article-title":"Attenuation of Japanese encephalitis virus by selection of its\u00a0mouse brain membrane receptor preparation escape variants","volume":"241","author":"Ni","year":"1998","journal-title":"Virology"},{"key":"10.1016\/j.heliyon.2019.e02882_bib26","doi-asserted-by":"crossref","first-page":"11467","DOI":"10.1128\/JVI.01125-06","article-title":"Crystal structure of the West Nile virus envelope glycoprotein","volume":"80","author":"Nybakken","year":"2006","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib27","series-title":"Fields Virology","first-page":"747","article-title":"Flaviviruses","author":"Pierson","year":"2013"},{"key":"10.1016\/j.heliyon.2019.e02882_bib28","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1038\/375291a0","article-title":"The envelope glycoprotein from tick-borne encephalitis virus at 2 A resolution","volume":"375","author":"Rey","year":"1995","journal-title":"Nature"},{"key":"10.1016\/j.heliyon.2019.e02882_bib29","doi-asserted-by":"crossref","first-page":"4522","DOI":"10.1128\/JVI.02686-13","article-title":"Dynamics of the emergence and establishment of a newly dominant genotype of Japanese encephalitis virus throughout Asia","volume":"88","author":"Schuh","year":"2014","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib30","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.jviromet.2005.02.001","article-title":"Detection of West Nile virus and Japanese encephalitis virus using real-time PCR with a probe common to both viruses","volume":"126","author":"Shirato","year":"2005","journal-title":"J.\u00a0Virol. Methods"},{"key":"10.1016\/j.heliyon.2019.e02882_bib31","doi-asserted-by":"crossref","first-page":"3091","DOI":"10.1128\/JVI.77.5.3091-3098.2003","article-title":"Origin and evolution of Japanese encephalitis virus in southeast Asia","volume":"77","author":"Solomon","year":"2003","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib32","doi-asserted-by":"crossref","first-page":"1144","DOI":"10.1093\/infdis\/171.5.1144","article-title":"Characterization of a highly attenuated Japanese encephalitis virus generated from molecularly cloned cDNA","volume":"171","author":"Sumiyoshi","year":"1995","journal-title":"J.\u00a0Infect. Dis."},{"key":"10.1016\/j.heliyon.2019.e02882_bib33","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.virol.2009.10.035","article-title":"A\u00a0single mutation in the Japanese encephalitis virus E protein (S123R) increases its growth rate in mouse neuroblastoma cells and its pathogenicity in mice","volume":"396","author":"Tajima","year":"2010","journal-title":"Virology"},{"key":"10.1016\/j.heliyon.2019.e02882_bib34","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.virusres.2005.08.015","article-title":"Nineteen nucleotides in the variable region of 3' non-translated region are dispensable for the replication of dengue type 1 virus in vitro","volume":"116","author":"Tajima","year":"2006","journal-title":"Virus Res."},{"key":"10.1016\/j.heliyon.2019.e02882_bib35","doi-asserted-by":"crossref","first-page":"2661","DOI":"10.1099\/vir.0.000213","article-title":"In vitro growth, pathogenicity and serological characteristics of the Japanese encephalitis virus genotype V Muar strain","volume":"96","author":"Tajima","year":"2015","journal-title":"J.\u00a0Gen. Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib36","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1186\/1743-422X-8-449","article-title":"Emergence of Japanese encephalitis virus genotype V in the Republic of Korea","volume":"8","author":"Takhampunya","year":"2011","journal-title":"Virol. J."},{"issue":"Suppl 2","key":"10.1016\/j.heliyon.2019.e02882_bib37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0264-410X(00)00037-2","article-title":"New initiatives for the control of Japanese encephalitis by vaccination: minutes of a WHO\/CVI meeting, Bangkok, Thailand, 13-15 October 1998","volume":"18","author":"Tsai","year":"2000","journal-title":"Vaccine"},{"key":"10.1016\/j.heliyon.2019.e02882_bib38","doi-asserted-by":"crossref","first-page":"242","DOI":"10.4269\/ajtmh.2001.65.242","article-title":"Phylogenetic analysis of Japanese encephalitis virus: envelope gene based analysis reveals a fifth genotype, geographic clustering, and multiple introductions of the virus into the Indian subcontinent","volume":"65","author":"Uchil","year":"2001","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"10.1016\/j.heliyon.2019.e02882_bib39","doi-asserted-by":"crossref","first-page":"46007","DOI":"10.1074\/jbc.M307776200","article-title":"Structural basis of a flavivirus recognized by its neutralizing antibody: solution structure of the domain III of the Japanese encephalitis virus envelope protein","volume":"278","author":"Wu","year":"2003","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.heliyon.2019.e02882_bib40","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/S0168-1702(00)00235-5","article-title":"Complete nucleotide sequence and cell-line multiplication pattern of the attenuated variant CH2195LA of Japanese encephalitis virus","volume":"73","author":"Wu","year":"2001","journal-title":"Virus Res."},{"key":"10.1016\/j.heliyon.2019.e02882_bib41","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1099\/vir.0.044925-0","article-title":"Characterization of a serine-to-asparagine substitution at position 123 in the Japanese encephalitis virus E protein","volume":"94","author":"Yamaguchi","year":"2013","journal-title":"J.\u00a0Gen. Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib42","series-title":"Envelope Protein Mutations L107F and E138K Are Important for Neurovirulence Attenuation for Japanese Encephalitis Virus SA14-14-2 Strain","author":"Yang","year":"2017"},{"key":"10.1016\/j.heliyon.2019.e02882_bib43","doi-asserted-by":"crossref","first-page":"3635","DOI":"10.1016\/j.vaccine.2010.02.105","article-title":"Phenotypic and genotypic characteristics of Japanese encephalitis attenuated live vaccine virus SA14-14-2 and their stabilities","volume":"28","author":"Yu","year":"2010","journal-title":"Vaccine"},{"key":"10.1016\/j.heliyon.2019.e02882_bib44","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1126\/science.aam7120","article-title":"A\u00a0single mutation in the prM protein of Zika virus contributes to fetal microcephaly","volume":"358","author":"Yuan","year":"2017","journal-title":"Science"},{"key":"10.1016\/j.heliyon.2019.e02882_bib45","doi-asserted-by":"crossref","first-page":"2209","DOI":"10.1099\/vir.0.80638-0","article-title":"Characterization of the E-138 (Glu\/Lys) mutation in Japanese encephalitis virus by using a stable, full-length, infectious cDNA clone","volume":"86","author":"Zhao","year":"2005","journal-title":"J.\u00a0Gen. Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib46","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.00108-18","article-title":"Acidity\/alkalinity of Japanese encephalitis virus E protein residue 138 alters neurovirulence in mice","volume":"92","author":"Zheng","year":"2018","journal-title":"J.\u00a0Virol."},{"key":"10.1016\/j.heliyon.2019.e02882_bib47","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1007\/s00705-018-3765-2","article-title":"Mutation of I176R in the E coding region weakens Japanese encephalitis virus neurovirulence, but not its growth rate in BHK-21 cells","volume":"163","author":"Zhou","year":"2018","journal-title":"Arch. Virol."}],"container-title":["Heliyon"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405844019365417?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405844019365417?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,2,2]],"date-time":"2025-02-02T04:02:45Z","timestamp":1738468965000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2405844019365417"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,11]]},"references-count":47,"journal-issue":{"issue":"11","published-print":{"date-parts":[[2019,11]]}},"alternative-id":["S2405844019365417"],"URL":"https:\/\/doi.org\/10.1016\/j.heliyon.2019.e02882","relation":{},"ISSN":["2405-8440"],"issn-type":[{"value":"2405-8440","type":"print"}],"subject":[],"published":{"date-parts":[[2019,11]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"E and prM proteins of genotype V Japanese encephalitis virus are required for its increased virulence in mice","name":"articletitle","label":"Article Title"},{"value":"Heliyon","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.heliyon.2019.e02882","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2019 The Author(s). Published by Elsevier Ltd.","name":"copyright","label":"Copyright"}],"article-number":"e02882"}}