{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,27]],"date-time":"2026-09-27T17:33:27Z","timestamp":1790530407709,"version":"4.1.0"},"reference-count":57,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2019,1,6]],"date-time":"2019-01-06T00:00:00Z","timestamp":1546732800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>Japanese encephalitis virus (JEV) is a flavivirus that is maintained via transmission between Culex spp. mosquitoes and water birds across a large swath of southern Asia and northern Australia. Currently JEV is the leading cause of vaccine-preventable encephalitis in humans in Asia. Five genotypes of JEV (G-I\u2013G-V) have been responsible for historical and current outbreaks in endemic regions, and G-I and G-III co-circulate throughout Southern Asia. While G-III has historically been the dominant genotype worldwide, G-I has gradually but steadily displaced G-III. The objective of this study was to better understand the phenomenon of genotype displacement for JEV by evaluating both avian host and mosquito vector susceptibilities to infection with representatives from both G-I and G-III. Since ducks and Culex quinquefasciatus mosquitoes are prevalent avian hosts and vectors perpetuating JEV transmission in JE endemic areas, experimental evaluation of virus replication in these species was considered to approximate the natural conditions necessary for studying the role of host, vectors and viral fitness in the JEV genotype displacement context. We evaluated viremia in ducklings infected with G-I and G-III, and did not detect differences in magnitude or duration of viremia. Testing the same viruses in mosquitoes revealed that the rates of infection, dissemination and transmission were higher in virus strains belonging to G-I than G-III, and that the extrinsic incubation period was shorter for the G-I strains. These data suggest that the characteristics of JEV infection of mosquitoes but not of ducklings, may have play a role in genotype displacement.<\/jats:p>","DOI":"10.3390\/v11010032","type":"journal-article","created":{"date-parts":[[2019,1,9]],"date-time":"2019-01-09T03:06:06Z","timestamp":1547003166000},"page":"32","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":31,"title":["Experimental Evaluation of the Role of Ecologically-Relevant Hosts and Vectors in Japanese Encephalitis Virus Genotype Displacement"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5557-4402","authenticated-orcid":false,"given":"Ajit K.","family":"Karna","sequence":"first","affiliation":[{"name":"Department of Microbiology, Immunology, and Pathology, Colorado State University, Fort Collins, CO 80523, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard A.","family":"Bowen","sequence":"additional","affiliation":[{"name":"Department of Microbiology, Immunology, and Pathology, Colorado State University, Fort Collins, CO 80523, USA"},{"name":"Department of Biomedical Sciences, Colorado State University, Fort Collins, CO 80523, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2019,1,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1146\/annurev.mi.44.100190.003245","article-title":"Flavivirus genome organization, expression, and replication","volume":"44","author":"Chambers","year":"1990","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/S0065-3527(03)59002-9","article-title":"Molecular biology of flaviviruses","volume":"59","author":"Lindenbach","year":"2003","journal-title":"Adv. Virus Res."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Lord, J.S., Gurley, E.S., and Pulliam, J.R. (2015). Rethinking Japanese Encephalitis Virus Transmission: A Framework for Implicating Host and Vector Species. PLoS Negl. Trop. Dis., 9.","DOI":"10.1371\/journal.pntd.0004074"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"S98","DOI":"10.1038\/nm1144","article-title":"Emerging flaviviruses: The spread and resurgence of Japanese encephalitis, West Nile and dengue viruses","volume":"10","author":"Mackenzie","year":"2004","journal-title":"Nat. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1146\/annurev.ento.54.110807.090510","article-title":"Ecology and geographical expansion of Japanese encephalitis virus","volume":"54","author":"Ritchie","year":"2009","journal-title":"Annu. Rev. Entomol."},{"key":"ref_6","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. Virol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"3722","DOI":"10.1016\/j.vaccine.2004.03.027","article-title":"Protection against Japanese encephalitis virus strains representing four genotypes by passive transfer of sera raised against ChimeriVax-JE experimental vaccine","volume":"22","author":"Beasley","year":"2004","journal-title":"Vaccine"},{"key":"ref_8","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":"ref_9","doi-asserted-by":"crossref","first-page":"747","DOI":"10.4269\/ajtmh.2001.65.747","article-title":"The appearance of a second genotype of Japanese encephalitis virus in the Australasian region","volume":"65","author":"Pyke","year":"2001","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Jeffries, C.L., and Walker, T. (2015). The Potential Use of Wolbachia-Based Mosquito Biocontrol Strategies for Japanese Encephalitis. PLoS Negl. Trop. Dis., 9.","DOI":"10.1371\/journal.pntd.0003576"},{"key":"ref_11","first-page":"1","article-title":"Japanese encephalitis vaccines: Recommendations of the Advisory Committee on Immunization Practices (ACIP)","volume":"59","author":"Fischer","year":"2010","journal-title":"MMWR Recomm. Rep."},{"key":"ref_12","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":"ref_13","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/s12250-015-3621-4","article-title":"Investigation of the genotype III to genotype I shift in Japanese encephalitis virus and the impact on human cases","volume":"30","author":"Han","year":"2015","journal-title":"Virol. Sin."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"2915","DOI":"10.1099\/0022-1317-71-12-2915","article-title":"Genetic variation of Japanese encephalitis virus in nature","volume":"71","author":"Chen","year":"1990","journal-title":"J. Gen. Virol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1016\/0168-1702(96)01306-8","article-title":"Phylogenetic analysis of the envelope gene of Japanese encephalitis virus","volume":"42","author":"Paranjpe","year":"1996","journal-title":"Virus Res."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2471","DOI":"10.1099\/0022-1317-81-10-2471","article-title":"Molecular characterization of the first Australian isolate of Japanese encephalitis virus, the FU strain","volume":"81","author":"Williams","year":"2000","journal-title":"J. Gen. Virol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Schuh, A.J., Ward, M.J., Brown, A.J., and Barrett, A.D. (2013). Phylogeography of Japanese encephalitis virus: Genotype is associated with climate. PLoS Negl. Trop. Dis., 7.","DOI":"10.1371\/journal.pntd.0002411"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1111\/j.1348-0421.1997.tb01196.x","article-title":"Antigenic and genetic variations among Japanese encephalitis virus strains belonging to genotype 1","volume":"41","author":"Ali","year":"1997","journal-title":"Microbiol. Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1016\/0264-410X(93)90262-V","article-title":"Immunogenicity of wild-type and vaccine strains of Japanese encephalitis virus and the effect of haplotype restriction on murine immune responses","volume":"11","author":"Wills","year":"1993","journal-title":"Vaccine"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.vaccine.2013.10.055","article-title":"Cross-protection elicited by primary and booster vaccinations against Japanese encephalitis: A two-year follow-up study","volume":"32","author":"Erra","year":"2013","journal-title":"Vaccine"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Bonaparte, M., Dweik, B., Feroldi, E., Meric, C., Bouckenooghe, A., Hildreth, S., Hu, B., Yoksan, S., and Boaz, M. (2014). Immune response to live-attenuated Japanese encephalitis vaccine (JE-CV) neutralizes Japanese encephalitis virus isolates from South-East Asia and India. BMC Infect. Dis., 14.","DOI":"10.1186\/1471-2334-14-156"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2165","DOI":"10.1099\/vir.0.000160","article-title":"Genotype-specific neutralization determinants in envelope protein: Implications for the improvement of Japanese encephalitis vaccine","volume":"96","author":"Ye","year":"2015","journal-title":"J. Gen. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"10832","DOI":"10.1038\/ncomms10832","article-title":"Vector-free transmission and persistence of Japanese encephalitis virus in pigs","volume":"7","author":"Ricklin","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"151","DOI":"10.4269\/ajtmh.2003.69.151","article-title":"Short report: A major genotype of Japanese encephalitis virus currently circulating in Japan","volume":"69","author":"Ma","year":"2003","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1099\/vir.0.79797-0","article-title":"Shift in Japanese encephalitis virus (JEV) genotype circulating in northern Vietnam: Implications for frequent introductions of JEV from Southeast Asia to East Asia","volume":"85 Pt 6","author":"Nga","year":"2004","journal-title":"J. Gen. Virol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1016\/j.meegid.2011.01.020","article-title":"Molecular phylogenetic and evolutionary analyses of Muar strain of Japanese encephalitis virus reveal it is the missing fifth genotype","volume":"11","author":"Mohammed","year":"2011","journal-title":"Infect. Genet. Evol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1186\/1743-422X-9-271","article-title":"Molecular evidence for the occurrence of Japanese encephalitis virus genotype I and III infection associated with acute encephalitis in patients of West Bengal, India, 2010","volume":"9","author":"Sarkar","year":"2012","journal-title":"Virol. J."},{"key":"ref_28","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. Virol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2354","DOI":"10.3201\/eid1712.110914","article-title":"Japanese encephalitis virus genotype replacement, Taiwan, 2009\u20132010","volume":"17","author":"Chen","year":"2011","journal-title":"Emerg. Infect. Dis."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"9847","DOI":"10.1128\/JVI.00825-11","article-title":"Emergence of genotype I of Japanese encephalitis virus as the dominant genotype in Asia","volume":"85","author":"Pan","year":"2011","journal-title":"J. Virol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1883","DOI":"10.3201\/eid2311.161443","article-title":"Virulence of Japanese Encephalitis Virus Genotypes I and III, Taiwan","volume":"23","author":"Fan","year":"2017","journal-title":"Emerg. Infect. Dis."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"493","DOI":"10.4269\/ajtmh.2004.71.493","article-title":"Genetic and phenotypic variation of West Nile virus in New York, 2000\u20132003","volume":"71","author":"Ebel","year":"2004","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"365","DOI":"10.4269\/ajtmh.2007.77.365","article-title":"A newly emergent genotype of West Nile virus is transmitted earlier and more efficiently by Culex mosquitoes","volume":"77","author":"Moudy","year":"2007","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1089\/153036601316977769","article-title":"Differential susceptibility of Aedes aegypti to infection by the American and Southeast Asian genotypes of dengue type 2 virus","volume":"1","author":"Armstrong","year":"2001","journal-title":"Vector Borne Zoonotic Dis."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Hanley, K.A., Nelson, J.T., Schirtzinger, E.E., Whitehead, S.S., and Hanson, C.T. (2008). Superior infectivity for mosquito vectors contributes to competitive displacement among strains of dengue virus. BMC Ecol., 8.","DOI":"10.1186\/1472-6785-8-1"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"13125","DOI":"10.1128\/JVI.01822-14","article-title":"Increased replicative fitness of a dengue virus 2 clade in native mosquitoes: Potential contribution to a clade replacement event in Nicaragua","volume":"88","author":"Quiner","year":"2014","journal-title":"J. Virol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1186\/s12985-015-0278-4","article-title":"Molecular epidemiology of Japanese encephalitis in northern Vietnam, 1964\u20132011: Genotype replacement","volume":"12","author":"Do","year":"2015","journal-title":"Virol. J."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"827","DOI":"10.1099\/vir.0.007617-0","article-title":"Evidence of frequent introductions of Japanese encephalitis virus from south-east Asia and continental east Asia to Japan","volume":"90","author":"Nabeshima","year":"2009","journal-title":"J. Gen. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"876","DOI":"10.3201\/eid1605.091055","article-title":"Molecular epidemiology of Japanese encephalitis virus, Taiwan","volume":"16","author":"Huang","year":"2010","journal-title":"Emerg. Infect. Dis."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Huang, Y.S., Hettenbach, S.M., Park, S.L., Higgs, S., Barrett, A.D., Hsu, W.W., Harbin, J.N., Cohnstaedt, L.W., and Vanlandingham, D.L. (2016). Differential Infectivities among Different Japanese Encephalitis Virus Genotypes in Culex quinquefasciatus Mosquitoes. PLoS Negl. Trop. Dis., 10.","DOI":"10.1371\/journal.pntd.0005038"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"242","DOI":"10.4269\/ajtmh.13-0161","article-title":"Age-related susceptibility to Japanese encephalitis virus in domestic ducklings and chicks","volume":"90","author":"Cleton","year":"2014","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1146\/annurev.mi.40.100186.002143","article-title":"The natural history of Japanese encephalitis virus","volume":"40","author":"Rosen","year":"1986","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1196\/annals.1373.013","article-title":"A serological survey of pigs, horses, and ducks in Nepal for evidence of infection with Japanese encephalitis virus","volume":"1081","author":"Pant","year":"2006","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1016\/0035-9203(70)90070-2","article-title":"Japanese encephalitis in Sarawak: Virus isolation and serology in a Land Dyak village","volume":"64","author":"Simpson","year":"1970","journal-title":"Trans. R. Soc. Trop. Med. Hyg."},{"key":"ref_45","first-page":"881","article-title":"Experimental viraemia and transmission of Japanese encephalitis virus by mosquitoes in domestic ducks","volume":"66","author":"Dhanda","year":"1977","journal-title":"Indian J. Med. Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"4703","DOI":"10.3390\/v6114703","article-title":"Flavivirus-mosquito interactions","volume":"6","author":"Huang","year":"2014","journal-title":"Viruses"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"482","DOI":"10.4269\/ajtmh.1972.21.482","article-title":"Experimental transmission of Japanese encephalitis virus by Culex fuscocephala","volume":"21","author":"Muangman","year":"1972","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_48","first-page":"875","article-title":"First isolation of Japanese encephalitis from Culex quinquefasciatus in Thailand","volume":"36","author":"Nitatpattana","year":"2005","journal-title":"Southeast. Asian J. Trop. Med. Public Health"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1126\/science.1122438","article-title":"Global patterns of influenza a virus in wild birds","volume":"312","author":"Olsen","year":"2006","journal-title":"Science"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1016\/j.agee.2006.09.001","article-title":"Avian influenza, domestic ducks and rice agriculture in Thailand","volume":"119","author":"Gilbert","year":"2007","journal-title":"Agric. Ecosyst. Environ."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"4769","DOI":"10.1073\/pnas.0710581105","article-title":"Mapping H5N1 highly pathogenic avian influenza risk in Southeast Asia","volume":"105","author":"Gilbert","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1080\/01652176.2017.1298885","article-title":"Duck virus enteritis (duck plague)\u2014A comprehensive update","volume":"37","author":"Dhama","year":"2017","journal-title":"Vet. Q."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Miller, R.H., Masuoka, P., Klein, T.A., Kim, H.C., Somer, T., and Grieco, J. (2012). Ecological niche modeling to estimate the distribution of Japanese encephalitis virus in Asia. PLoS Negl. Trop. Dis., 6.","DOI":"10.1371\/journal.pntd.0001678"},{"key":"ref_54","first-page":"709","article-title":"Experimental viraemia and transmission of Japanese encephalitis virus by mosquitoes in ardeid birds","volume":"66","author":"Soman","year":"1977","journal-title":"Indian J. Med. Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/S0168-7069(06)16010-3","article-title":"Japanese encephalitis virus: The geographic distribution, incidence, and spread of a virus with a propensity to emerge in new areas","volume":"16","author":"Mackenzie","year":"2006","journal-title":"Perspect. Med. Virol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"690","DOI":"10.4269\/ajtmh.1984.33.690","article-title":"The effect of colonization upon aedes aegypti susceptibility to oral infection with yellow fever virus","volume":"33","author":"Lorenz","year":"1984","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"2704","DOI":"10.4161\/hv.29509","article-title":"Reduction of neutralization antibody against heterologous circulating strains in adults immunized with Japanese encephalitis live vaccine","volume":"10","author":"Ye","year":"2014","journal-title":"Hum. Vaccin Immunother."}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/11\/1\/32\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T12:23:51Z","timestamp":1760185431000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/11\/1\/32"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,1,6]]},"references-count":57,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,1]]}},"alternative-id":["v11010032"],"URL":"https:\/\/doi.org\/10.3390\/v11010032","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,1,6]]}}}