Japanese encephalitis virus (JEV) is a reemerging vector-borne orthoflavivirus of significant public and animal health concern. Infection with JEV can cause inapparent, mild, or fatal encephalitis in humans and animals. The virus, which circulates primarily in Southeast Asia, has caused outbreaks in humans and livestock in nonendemic regions, leading to human disease and significant economic loss. The 2022 Australian outbreak of JEV led to renewed interest in JEV countermeasures available in the United States for veterinary use as the Australia outbreak primarily affected commercial swine farms. There are currently no antivirals or veterinary vaccines available to protect animals from JEV infection, and only one vaccine is approved for human use in the United States. In the event of a JEV introduction to the United States, vaccination of susceptible reservoirs, like domestic swine, may help prevent economic loss and JEV from establishing endemicity in the United States.
Methods:
This review assesses recent advances in JEV vaccine development with an interest in veterinary applications. This systematic review was conducted by searching databases (PubMed, Web of Science, Scopus, and EBSCOhost).
Results:
Eighty-seven research articles focused on the evaluation of novel JEV vaccine candidates were included in this review.
Conclusion:
Although there are a variety of different methodologies being employed in the development of JEV vaccine candidates, including recombinant live virus vaccines utilizing insect-only flaviviruses as vectors for JEV antigens and virus-like particle vaccines using various JEV genes to elicit protection among different JEV genotypes. While there are many promising candidates being developed, most are likely several years away from commercial production and would not be available if JEV were introduced to the United States within the next several years. Therefore, other disease control strategies should be investigated.
AdugnaT, NiuQ, GuanG, et al.Advancements in nanoparticle-based vaccine development against Japanese encephalitis virus: A systematic review. Front Immunol, 2024; 15:1505612.
2.
AhnJ, YuJE, KimH, et al.AB(5)-type toxin as a pentameric scaffold in recombinant vaccines against the Japanese encephalitis virus. Toxins (Basel), 2023; 15(7):425.
3.
AhsanMF, GoreMM. Comparison of immune response generated against Japanese encephalitis virus envelope protein expressed by DNA vaccines under macrophage associated versus ubiquitous expression promoters. Virol J, 2011; 8:382.
4.
AnwarMN, JiangCY, DiD, et al.A novel recombinant virus-like particles displaying B and T cell epitopes of Japanese encephalitis virus offers protective immunity in mice and guinea pigs. Vaccines (Basel), 2021; 9(9):980.
5.
ArakawaT, HarakuniT, MiyataT, et al.Tricomponent fusion complex comprising a viral antigen, a pentameric alpha-helical coiled-coil, and an immunoglobulin-binding domain as an effective antiviral vaccine. Vaccine, 2014; 32(7):864–871.
6.
BelloMB, AlsaadiA, NaeemA, et al.Development of nucleic acid-based vaccines against dengue and other mosquito-borne flaviviruses: The past, present, and future. Front Immunol, 2024; 15:1475886.
7.
Bielefeldt-OhmannH, ProwNA, WangW, et al.Safety and immunogenicity of a delta inulin-adjuvanted inactivated Japanese encephalitis virus vaccine in pregnant mares and foals. Vet Res, 2014; 45(1):130.
ChangYH, ChiaoDJ, HsuYL, et al.Mosquito cell-derived Japanese encephalitis virus-like particles induce specific humoral and cellular immune responses in mice. Viruses-Basel, 2020; 12(3):336.
10.
ChenH, GaoN, FanD, et al.Suppressive effects on the immune response and protective immunity to a JEV DNA vaccine by co-administration of a GM-CSF-expressing plasmid in mice. PLoS One, 2012; 7(4):e34602.
11.
ChenT, ZhuS, WeiN, et al.Protective immune responses induced by an mRNA-LNP vaccine encoding prM-E proteins against Japanese encephalitis virus infection. Viruses, 2022; 14(6):1121.
12.
ChenTH, HuCC, LiaoJT, et al.Production of Japanese encephalitis virus antigens in plants using bamboo mosaic virus-based vector. Front Microbiol, 2017; 8:788.
13.
ChongHY, LeowCY, Abdul MajeedAB, et al.Flavivirus infection-A review of immunopathogenesis, immunological response, and immunodiagnosis. Virus Res, 2019; 274:197770.
de WispelaereM, KhouC, FrenkielMP, et al.A single amino acid substitution in the M protein attenuates Japanese encephalitis virus in mammalian hosts. J Virol, 2015a;90(5):2676–2689.
16.
de WispelaereM, RicklinM, SouqueP, et al.A lentiviral vector expressing Japanese encephalitis virus-like particles elicits broad neutralizing antibody response in pigs. PLoS Negl Trop Dis, 2015b;9(10):e0004081.
17.
DixonAL, OliveiraARS, CohnstaedtLW, et al.Revisiting the risk of introduction of Japanese encephalitis virus (JEV) into the United States—An updated semi-quantitative risk assessment. One Health, 2024; 19:100879.
18.
FanJ, LiuY, XieX, et al.Inhibition of Japanese encephalitis virus infection by flavivirus recombinant E protein domain III. Virol Sin, 2013a;28(3):152–160.
19.
FanYC, ChenJM, ChenYY, et al.Reduced neutralizing antibody titer against genotype I virus in swine immunized with a live-attenuated genotype III Japanese encephalitis virus vaccine. Vet Microbiol, 2013b;163(3–4):248–256.
20.
FanYC, ChenJM, LinJW, et al.Genotype I of Japanese encephalitis virus virus-like particles elicit sterilizing immunity against genotype I and III viral challenge in swine. Sci Rep, 2018; 8(1):7481.
21.
FanYC, ChenYY, ChenJM, et al.Effectiveness of live-attenuated genotype III Japanese encephalitis viral vaccine against circulating genotype I viruses in swine. Viruses, 2022; 14(1):114.
22.
FirbasC, JilmaB. Product review on the JE vaccine IXIARO. Hum Vaccin Immunother, 2015; 11(2):411–420.
23.
FujisakiY, SugimoriT, MorimotoT, et al.Immunization of pigs with the attenuated S-strain of Japanese encephalitis virus. Natl Inst Anim Health Q (Tokyo), 1975; 15(2):55–60.
24.
GangwarRS, ShilP, SapkalGN, et al.Induction of virus-specific neutralizing immune response against West Nile and Japanese encephalitis viruses by chimeric peptides representing T-helper and B-cell epitopes. Virus Res, 2012; 163(1):40–50.
25.
GaoN, ChenW, ZhengQ, et al.Co-expression of Japanese encephalitis virus prM-E-NS1 antigen with granulocyte-macrophage colony-stimulating factor enhances humoral and anti-virus immunity after DNA vaccination. Immunol Lett, 2010; 129(1):23–31.
26.
Garcia-NicolasO, RicklinME, LinigerM, et al.A Japanese encephalitis virus vaccine inducing antibodies strongly enhancing in vitro infection is protective in pigs. Viruses, 2017; 9(5):124.
27.
GargH, Mehmetoglu-GurbuzT, JoshiA. Virus like particles (VLP) as multivalent vaccine candidate against Chikungunya, Japanese encephalitis, yellow fever and zika virus. Sci Rep, 2020; 10(1):4017.
28.
GromowskiGD, FirestoneCY, Bustos-ArriagaJ, et al.Genetic and phenotypic properties of vero cell-adapted Japanese encephalitis virus SA14-14-2 vaccine strain variants and a recombinant clone, which demonstrates attenuation and immunogenicity in mice. Am J Trop Med Hyg, 2015; 92(1):98–107.
29.
GromowskiGD, FirestoneCY, HansonCT, et al.Japanese encephalitis virus vaccine candidates generated by chimerization with dengue virus type 4. Vaccine, 2014; 32(25):3010–3018.
30.
HalsteadSB, ThomasSJ. New Japanese encephalitis vaccines: Alternatives to production in mouse brain. Expert Rev Vaccines, 2011; 10(3):355–364.
31.
HarrisonJJ, NguyenW, MorganMS, et al.A chimeric vaccine derived from Australian genotype IV Japanese encephalitis virus protects mice from lethal challenge. NPJ Vaccines, 2024; 9(1):134.
32.
HiguchiA, ToriniwaH, KomiyaT, et al.Recombinant measles AIK-C vaccine strain expressing the prM-E antigen of Japanese encephalitis virus. PLoS One, 2016; 11(3):e0150213.
33.
HombachJ, SolomonT, KuraneI, et al.Report on a WHO consultation on immunological endpoints for evaluation of new Japanese encephalitis vaccines, WHO, Geneva, 2-3 September, 2004. Vaccine, 2005; 23(45):5205–5211.
34.
HonjoS, MasudaM, IshikawaT. Effects of the Japanese encephalitis virus genotype V-derived sub-viral particles on the immunogenicity of the vaccine characterized by a novel virus-like particle-based assay. Vaccines (Basel), 2019; 7(3):81.
35.
HsuST, ChangLC, LinSY, et al.The effect of vaccination with a live attenuated strain of Japanese encephalitis virus on stillbirths in swine in Taiwan. Bull World Health Organ, 1972; 46(4):465–471.
36.
HuaRH, LiYN, ChenZS, et al.Generation and characterization of a new mammalian cell line continuously expressing virus-like particles of Japanese encephalitis virus for a subunit vaccine candidate. BMC Biotechnol, 2014; 14:62.
37.
HuangY, LiuS, YangP, et al.Replicon-based Japanese encephalitis virus vaccines elicit immune response in mice. J Virol Methods, 2012; 179(1):217–225.
38.
HuangY, XuH, LiuS, et al.Accidental acquisition of a rescued Japanese encephalitis virus with unspliced introns in the viral genome when using an intron-based stabilization approach. Arch Virol, 2023; 168(2):60.
39.
HuangYS, HettenbachSM, ParkSL, et al.Differential infectivities among different Japanese encephalitis virus genotypes in Culex quinquefasciatus mosquitoes. PLoS Negl Trop Dis, 2016; 10(10):e0005038.
ImotoJ, IshikawaT, YamanakaA, et al.Needle-free jet injection of small doses of Japanese encephalitis DNA and inactivated vaccine mixture induces neutralizing antibodies in miniature pigs and protects against fetal death and mummification in pregnant sows. Vaccine, 2010; 28(46):7373–7380.
42.
IshikawaT, WangG, WidmanDG, et al.Enhancing the utility of a prM/E-expressing chimeric vaccine for Japanese encephalitis by addition of the JEV NS1 gene. Vaccine, 2011; 29(43):7444–7455.
43.
IslamMD, IslamMM, InoueA, et al.Neutralizing antibodies against the Japanese encephalitis virus are produced by a 12 kDa E. coli-expressed envelope protein domain III (EDIII) tagged with a solubility-controlling peptide. Vaccine, 2025; 56:127143.
44.
JungJW, ParkPG, LeeWK, et al.Production of plant-derived Japanese encephalitis virus multi-epitope peptide in nicotiana benthamiana and immunological response in mice. Int J Mol Sci, 2023; 24(14):11643.
45.
KimHH, YangDK, JiM, et al.Genotype-specific neutralizing antibody titers against Japanese encephalitis virus genotypes 1 and 3 in horses immunized with a genotype 3 vaccine. Clin Exp Vaccine Res, 2020; 9(2):102–107.
46.
KimJD, LeeAR, MoonDH, et al.Efficacy of genotype-matched vaccine against re-emerging genotype V Japanese encephalitis virus. Emerg Microbes Infect, 2024; 13(1):2343910.
47.
KomiyaT, Honda-OkuboY, BaldwinJ, et al.An advax-adjuvanted inactivated cell-culture derived Japanese encephalitis vaccine induces broadly neutralising anti-flavivirus antibodies, robust cellular immunity and provides single dose protection. Vaccines (Basel), 2021; 9(11):1235.
48.
KotakiT, NagaiY, YamanakaA, et al.Japanese encephalitis DNA vaccines with epitope modification reduce the induction of cross-reactive antibodies against dengue virus and antibody-dependent enhancement of dengue virus infection. Vaccines (Basel), 2022; 10(9):1411.
49.
KulkarniR, SapkalG, GoreM. Evaluation of Japanese encephalitis virus polytope DNA vaccine candidate in BALB/c mice. Virus Res, 2012a;170(1–2):118–125.
50.
KulkarniR, SapkalG, MahishiL, et al.Design and characterization of polytope construct with multiple B and TH epitopes of Japanese encephalitis virus. Virus Res, 2012b;166(1–2):77–86.
51.
KumarK, OngHK, TanWS, et al.Immunological analysis of nodavirus capsid displaying the domain III of Japanese encephalitis virus envelope protein. Pharmaceutics, 2021; 13(11):1826.
52.
KwonWT, LeeWS, ParkPJ, et al.Protective immunity of Pichia pastoris-expressed recombinant envelope protein of Japanese encephalitis virus. J Microbiol Biotechnol, 2012; 22(11):1580–1587.
53.
LeeE, KimM, KimYB. Attenuated chimeric GI/GIII vaccine candidate against Japanese encephalitis virus. Vaccines (Basel), 2023; 11(12):1827.
54.
LeeEY, KimJY, LeeDK, et al.Sublingual immunization with Japanese encephalitis virus vaccine effectively induces immunity through both cellular and humoral immune responses in mice. Microbiol Immunol, 2016; 60(12):846–853.
55.
LeeHH, HongSK, YoonSH, et al.Immunogenicity of Japanese encephalitis virus envelope protein by Hyphantria cunea nuclear polyhedrosis virus vector in guinea pig. Appl Biochem Biotechnol, 2012; 167(2):259–269.
56.
LiJ, ChenH, WuN, et al.Characterization of immune responses induced by inactivated, live attenuated and DNA vaccines against Japanese encephalitis virus in mice. Vaccine, 2013; 31(38):4136–4142.
57.
LiN, ZhangZR, ZhangYN, et al.A replication-defective Japanese encephalitis virus (JEV) vaccine candidate with NS1 deletion confers dual protection against JEV and West Nile virus in mice. NPJ Vaccines, 2020; 5(1):73.
58.
LiP, CaoRB, ZhengQS, et al.Enhancement of humoral and cellular immunity in mice against Japanese encephalitis virus using a DNA prime-protein boost vaccine strategy. Vet J, 2010; 183(2):210–216.
59.
LiZ, ZhangJ, LiH. Flavivirus NS2B/NS3 protease: Structure, function, and inhibition. In: Viral Proteases and Their Inhibitors. Academic Press; 2017; pp.163–188.
60.
LienSB, YangQW, HuangHW, et al.A novel immunogen comprising a bc loop and mutant fusion loop epitopes generates potent neutralization and protective abilities against flaviviruses without risk of disease enhancement. Vaccine, 2025; 57:127219.
61.
LinTS, ChuangCC, HsuHL, et al.Role of amphotericin B upon enhancement of protective immunity elicited by oral administration with liposome-encapsulated-Japanese encephalitis virus nonstructural protein 1 (NS1) in mice. Microb Pathog, 2010; 49(3):67–74.
62.
LindbladEB. Freund’s adjuvant. In: Vaccine Adjuvants: preparation methods and research protocols. (O’Hagan DT., ed.) Humana Press; 2000; pp. 49–64.
63.
LiuH, WuR, LiuK, et al.Enhanced immune responses against Japanese encephalitis virus using recombinant adenoviruses coexpressing Japanese encephalitis virus envelope and porcine interleukin-6 proteins in mice. Virus Res, 2016; 222:34–40.
64.
LiuX, YuY, LiM, et al.Study on the protective efficacy of SA14-14-2 attenuated Japanese encephalitis against different JE virus isolates circulating in China. Vaccine, 2011; 29(11):2127–2130.
65.
LobigsM, PavyM, HallRA, et al.An inactivated Vero cell-grown Japanese encephalitis vaccine formulated with Advax, a novel inulin-based adjuvant, induces protective neutralizing antibody against homologous and heterologous flaviviruses. J Gen Virol, 2010; 91(Pt 6):1407–1417.
LucaVC, AbiMansourJ, NelsonCA, et al.Crystal structure of the Japanese encephalitis virus envelope protein. J Virol, 2012; 86(4):2337–2346.
68.
MackenzieJS, BarrettAD, DeubelV. Japanese Encephalitis and West Nile Virues. Springer; 2002.
69.
MackenzieJS, WilliamsDT, van den HurkAF, et al.Japanese encephalitis virus: The emergence of genotype IV in Australia and its potential endemicity. Viruses, 2022; 14(11):2480.
70.
MansfieldKL, Hernandez-TrianaLM, BanyardAC, et al.Japanese encephalitis virus infection, diagnosis and control in domestic animals. Vet Microbiol, 2017; 201:85–92.
71.
MatsudaS, NeromeR, MaegawaK, et al.Development of a Japanese encephalitis virus-like particle vaccine in silkworms using codon-optimised prM and envelope genes. Heliyon, 2017; 3(4):e00286.
72.
MatsuoH, SomiyaM, IijimaM, et al.CD11c-specific bio-nanocapsule enhances vaccine immunogenicity by targeting immune cells. J Nanobiotechnology, 2018; 16(1):59.
73.
McArthurMA, HolbrookMR. Japanese encephalitis vaccines. J Bioterror Biodef, 2011; S1:2.
74.
MisraUK, KalitaJ. Overview: Japanese encephalitis. Prog Neurobiol, 2010; 91(2):108–120.
75.
MiyataT, TafukuS, HarakuniT, et al.A bio-nanocapsule containing envelope protein domain III of Japanese encephalitis virus protects mice against lethal Japanese encephalitis virus infection. Microbiol Immunol, 2013; 57(6):470–477.
MooreSM. The current burden of Japanese encephalitis and the estimated impacts of vaccination: Combining estimates of the spatial distribution and transmission intensity of a zoonotic pathogen. PLoS Negl Trop Dis, 2021; 15(10):e0009385.
78.
NahJJ, YangDK, KimHH, et al.The present and future of veterinary vaccines for Japanese encephalitis in Korea. Clin Exp Vaccine Res, 2015; 4(2):130–136.
79.
NathB, SainiHM, PrasadM, et al.Evaluation of Japanese encephalitis virus E and NS1 proteins immunogenicity using a recombinant Newcastle disease virus in mice. Vaccine, 2020; 38(7):1860–1868.
80.
NemethN, Bosco-LauthA, OesterleP, et al.North American birds as potential amplifying hosts of Japanese encephalitis virus. Am J Trop Med Hyg, 2012; 87(4):760–767.
81.
NeromeK, YamaguchiR, FukeN, et al.Development of a Japanese encephalitis virus genotype V virus-like particle vaccine in silkworms. J Gen Virol, 2018; 99(7):897–907.
82.
NickolsB, TretyakovaI, TibbensA, et al.Plasmid DNA launches live-attenuated Japanese encephalitis virus and elicits virus-neutralizing antibodies in BALB/c mice. Virology, 2017; 512:66–73.
83.
NielsenHS, OleksiewiczMB, ForsbergR, et al.Reversion of a live porcine reproductive and respiratory syndrome virus vaccine investigated by parallel mutations. J Gen Virol, 2001; 82(Pt 6):1263–1272.
84.
OkamotoS, YoshiiH, MatsuuraM, et al.Poly-gamma-glutamic acid nanoparticles and aluminum adjuvant used as an adjuvant with a single dose of Japanese encephalitis virus-like particles provide effective protection from Japanese encephalitis virus. Clin Vaccine Immunol, 2012; 19(1):17–22.
85.
OliveiraARS, CohnstaedtLW, NoronhaLE, et al.Perspectives regarding the risk of introduction of the Japanese encephalitis virus (JEV) in the United States. Front Vet Sci, 2020; 7:48.
86.
OliveiraARS, PiaggioJ, CohnstaedtLW, et al.A quantitative risk assessment (QRA) of the risk of introduction of the Japanese encephalitis virus (JEV) in the United States via infected mosquitoes transported in aircraft and cargo ships. Prev Vet Med, 2018; 160:1–9.
87.
OliveiraARS, PiaggioJ, CohnstaedtLW, et al.Introduction of the Japanese encephalitis virus (JEV) in the United States—A qualitative risk assessment. Transbound Emerg Dis, 2019; 66(4):1558–1574.
88.
ParkJY, LeeHM, JunSH, et al.Insights into the pathogenesis and development of recombinant Japanese encephalitis virus genotype 3 as a vaccine. Vaccines (Basel), 2024; 12(6):597.
89.
ParkSL, HuangYJS, LyonsAC, et al.Mosquito saliva modulates Japanese encephalitis virus infection in domestic pigs. FrontVirol, 2021; 1.
90.
ParkSL, HuangYS, LyonsAC, et al.North American domestic pigs are susceptible to experimental infection with Japanese encephalitis virus. Sci Rep, 2018; 8(1):7951.
91.
QianP, ZhiX, WangB, et al.Construction and immune efficacy of recombinant pseudorabies virus expressing PrM-E proteins of Japanese encephalitis virus genotype capital I, Ukrainian. Virol J, 2015; 12:214.
92.
QuanTM, ThaoTTN, DuyNM, et al.Estimates of the global burden of Japanese encephalitis and the impact of vaccination from 2000–2015. Elife, 2020; 9:e51027.
93.
RavaniniP, HuhtamoE, IlariaV, et al.Japanese encephalitis virus RNA detected in Culex pipiens mosquitoes in Italy. Euro Surveill, 2012; 17(28):20221.
94.
ReevesWC, HammonWM. Laboratory transmission of Japanese B encephalitis virus by seven species (three genera) of North American mosquitoes. J Exp Med, 1946; 83(3):185–194.
95.
RumyantsevAA, Giel-MoloneyM, LiuY, et al.Characterization of the RepliVax platform for replication-defective flavivirus vaccines. Vaccine, 2011; 29(32):5184–5194.
96.
SchuhAJ, WardMJ, Leigh BrownAJ, et al.Dynamics of the emergence and establishment of a newly dominant genotype of Japanese encephalitis virus throughout Asia. J Virol, 2014; 88(8):4522–4532.
97.
ShahanaPV, DasD, GontuA, et al.Efficient production of Tymovirus like particles displaying immunodominant epitopes of Japanese encephalitis virus envelope protein. Protein Expr Purif, 2015; 113:35–43.
98.
ShahanaPV, DasD, GontuA, et al.Enhancement of Japanese encephalitis virus DNA vaccine efficacy by using Mycobacterium avium paratuberculosis heat shock protein 65. Curr Trends Biotechnol Pharm, 2013; 7(4):904–913.
99.
ShengZ, GaoN, CuiX, et al.Electroporation enhances protective immune response of a DNA vaccine against Japanese encephalitis in mice and pigs. Vaccine, 2016; 34(47):5751–5757.
100.
SunE, ZhaoJ, XuQ, et al.Antibodies generated by immunization with the NS1 protein of West Nile virus confer partial protection against lethal Japanese encephalitis virus challenge. Vet Microbiol, 2013; 166(1–2):145–153.
101.
TafukuS, MiyataT, TadanoM, et al.Japanese encephalitis virus structural and nonstructural proteins expressed in Escherichia coli induce protective immunity in mice. Microbes Infect, 2012; 14(2):169–176.
102.
TajimaS, TaniguchiS, NakayamaE, et al.Immunogenicity and protective ability of genotype I-based recombinant Japanese encephalitis virus (JEV) with attenuation mutations in E protein against genotype V JEV. Vaccines (Basel), 2021; 9(10):1077.
103.
ToriniwaH, KomiyaT. Comparison of viral glycosylation using lectin blotting with Vero cell-derived and mouse brain-derived Japanese encephalitis vaccines. Vaccine, 2011; 29(10):1859–1862.
104.
TripathiNK, ShrivastavaA, BiswalKC, et al.Development of a pilot-scale production process and characterization of a recombinant Japanese encephalitis virus envelope domain III protein expressed in Escherichia coli. Appl Microbiol Biotechnol, 2012; 95(5):1179–1189.
van den HurkAF, RitchieSA, MackenzieJS. Ecology and geographical expansion of Japanese encephalitis virus. Annu Rev Entomol, 2009; 54:17–35.
107.
WanJ, WangT, XuJ, et al.Novel Japanese encephalitis virus NS1-based vaccine: Truncated NS1 fused with E. coli heat labile enterotoxin B subunit. EBioMedicine, 2021; 67:103353.
108.
WangF, FengX, ZhengQ, et al.Multiple linear epitopes (B-cell, CTL and Th) of JEV expressed in recombinant MVA as multiple epitope vaccine induces a protective immune response. Virol J, 2012; 9:204.
109.
WangM, JiangS, WangY. Recent advances in the production of recombinant subunit vaccines in Pichia pastoris. Bioengineered, 2016; 7(3):155–165.
WongSH, JasseyA, WangJY, et al.Virus-like particle systems for vaccine development against viruses in the flaviviridae family. Vaccines (Basel), 2019; 7(4):123.
112.
XiaQ, YangY, ZhangY, et al.Shift in dominant genotypes of Japanese encephalitis virus and its impact on current vaccination strategies. Front Microbiol, 2023; 14:1302101.
113.
XiaQ, ZhangY, YangY, et al.Virulence and cross-protection conferred by an attenuated genotype I-based chimeric Japanese encephalitis virus strain harboring the E protein of genotype V in mice. Microbiol Spectr, 2022; 10(6):e0199022.
114.
XuB, LiuX, YanD, et al.Generation and characterization of chimeric Tembusu viruses containing pre-membrane and envelope genes of Japanese encephalitis virus. Front Microbiol, 2023; 14:1140141.
115.
XuXG, WangZS, ZhangQ, et al.Baculovirus surface display of E envelope glycoprotein of Japanese encephalitis virus and its immunogenicity of the displayed proteins in mouse and swine models. Vaccine, 2011; 29(4):636–643.
116.
XuY, WangQ, WeiB, et al.Enhanced immune responses against Japanese encephalitis virus infection using Japanese encephalitis live-attenuated virus adjuvanted with montanide GEL 01 ST in mice. Vector Borne Zoonotic Dis, 2019; 19(11):835–843.
117.
YamajiH, NakamuraM, KuwaharaM, et al.Efficient production of Japanese encephalitis virus-like particles by recombinant lepidopteran insect cells. Appl Microbiol Biotechnol, 2013; 97(3):1071–1079.
118.
YamanakaA, RattanaamnuaychaiP, MatsudaM, et al.Engineered flavivirus vaccines control induction of crossreactive infection-enhancing and -neutralizing antibodies. Vaccine, 2022; 40(42):6004–6011.
119.
YamanakaT, TsujimuraK, KondoT, et al.Isolation and genetic analysis of Japanese encephalitis virus from a diseased horse in Japan. J Vet Med Sci, 2006; 68(3):293–295.
120.
YangDK, NahJJ, KimHH, et al.Inactivated genotype 1 Japanese encephalitis vaccine for swine. Clin Exp Vaccine Res, 2014; 3(2):212–219.
121.
YangL, XiaoA, WangH, et al.A VLP-based vaccine candidate protects mice against Japanese encephalitis virus infection. Vaccines (Basel), 2022; 10(2):197.
122.
YangMH, HuCC, WongCH, et al.Convenient auto-processing vector based on bamboo mosaic virus for presentation of antigens through enzymatic coupling. Front Immunol, 2021; 12:739837.
123.
YangZ, WuR, LiRW, et al.Chimeric classical swine fever (CSF)-Japanese encephalitis (JE) viral replicon as a non-transmissible vaccine candidate against CSF and JE infections. Virus Res, 2012; 165(1):61–70.
124.
YaoM, RenX, YinM, et al.Nanoparticle vaccine based on the envelope protein domain III of Japanese encephalitis virus elicits robust protective immune responses in mice. Nanomedicine (Lond), 2023; 18(1):5–18.
125.
YenLC, ChenHW, HoCL, et al.Neutralizing antibodies targeting a novel epitope on envelope protein exhibited broad protection against flavivirus without risk of disease enhancement. J Biomed Sci, 2023; 30(1):41.
126.
ZangX, LiG, ZhuJ, et al.Evaluation of the adjuvant effect of imiquimod and CpG ODN 1826 in chimeric DNA vaccine against Japanese encephalitis. Int Immunopharmacol, 2024; 140:112816.
127.
ZhaiY, ZhouY, LiX, et al.Immune-enhancing effect of nano-DNA vaccine encoding a gene of the prME protein of Japanese encephalitis virus and BALB/c mouse granulocyte-macrophage colony-stimulating factor. Mol Med Rep, 2015; 12(1):199–209.
128.
ZhaiYZ, ZhouY, MaL, et al.The dominant roles of ICAM-1-encoding gene in DNA vaccination against Japanese encephalitis virus are the activation of dendritic cells and enhancement of cellular immunity. Cell Immunol, 2013; 281(1):1–10.
129.
ZhangH, ZhangY, LiD, et al.Partial protective efficacy of the current licensed Japanese encephalitis live vaccine against the emerging genotype I Japanese encephalitis virus isolated from sheep. Front Immunol, 2025; 16:1513261.
130.
ZhangY, ChenP, CaoR, et al.Mutation of putative N-linked glycosylation sites in Japanese encephalitis virus premembrane and envelope proteins enhances humoral immunity in BALB/C mice after DNA vaccination. Virol J, 2011; 8:138.
131.
ZhangY, HanJC, JingJ, et al.Construction and immunogenicity of recombinant vaccinia virus vaccine against Japanese encephalitis and chikungunya viruses infection in mice. Vector Borne Zoonotic Dis, 2020; 20(10):788–796.
132.
ZhaoF, ZhaiY, ZhuJ, et al.Enhancement of autophagy as a strategy for development of new DNA vaccine candidates against Japanese encephalitis. Vaccine, 2019; 37(37):5588–5595.
133.
ZhengX, YuX, WangY, et al.Immune responses and protective effects against Japanese encephalitis induced by a DNA vaccine encoding the prM/E proteins of the attenuated SA14-14-2 strain. Infect Genet Evol, 2020a;85:104443.
134.
ZhengX, YuX, WangY, et al.Complete protection for mice conferred by a DNA vaccine based on the Japanese encephalitis virus P3 strain used to prepare the inactivated vaccine in China. Virol J, 2020b;17(1):126.
135.
ZhuJ, HeC, LiuY, et al.An engineered Japanese encephalitis virus mRNA-lipid nanoparticle immunization induces protective immunity in mice. Front Microbiol, 2024; 15:1472824.
