{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T00:06:54Z","timestamp":1793232414433,"version":"4.5.0"},"reference-count":34,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2023,9,20]],"date-time":"2023-09-20T00:00:00Z","timestamp":1695168000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Natural Science Foundation of China","award":["32172823"],"award-info":[{"award-number":["32172823"]}]},{"name":"National Natural Science Foundation of China","award":["32102637"],"award-info":[{"award-number":["32102637"]}]},{"name":"National Natural Science Foundation of China","award":["SJCX22_1805"],"award-info":[{"award-number":["SJCX22_1805"]}]},{"name":"Postgraduate Research and Practice Innovation Program of Jiangsu Province (Yangzhou University)","award":["32172823"],"award-info":[{"award-number":["32172823"]}]},{"name":"Postgraduate Research and Practice Innovation Program of Jiangsu Province (Yangzhou University)","award":["32102637"],"award-info":[{"award-number":["32102637"]}]},{"name":"Postgraduate Research and Practice Innovation Program of Jiangsu Province (Yangzhou University)","award":["SJCX22_1805"],"award-info":[{"award-number":["SJCX22_1805"]}]},{"name":"Project of the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD)","award":["32172823"],"award-info":[{"award-number":["32172823"]}]},{"name":"Project of the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD)","award":["32102637"],"award-info":[{"award-number":["32102637"]}]},{"name":"Project of the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD)","award":["SJCX22_1805"],"award-info":[{"award-number":["SJCX22_1805"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>Pseudorabies virus (PRV) variants were discovered in immunized pigs in Northern China and have become the dominant strains since 2011, which caused huge economic losses. In this study, a classical PRV strain was successfully isolated in a PRV gE positive swine farm. The complete genome sequence was obtained using a high-throughput sequencing method and the virus was named JS-2020. The nucleotide homology analysis and phylogenetic tree based on complete genome sequences or gC gene showed that the JS-2020 strain was relatively close to the classical Ea strain in genotype II clade. However, a large number of amino acid variations occurred in the JS-2020 strain compared with the Ea strain, including multiple immunogenic and virulence-related genes. In particular, the gE protein of JS-2020 was similar to earlier Chinese PRV strains without Aspartate insertion. However, the amino acid variations analysis based on major immunogenic and virulence-related genes showed that the JS-2020 strain was not only homologous with earlier PRV strains, but also with strains isolated in recent years. Moreover, the JS-2020 strain was identified as a recombinant between the GXGG-2016 and HLJ-2013 strains. The pathogenicity analysis proved that the PRV JS-2020 strain has typical neurogenic infections and a strong pathogenicity in mice. Together, a novel recombinant classical strain was isolated and characterized in the context of the PRV variant pandemic in China. This study provided some valuable information for the study of the evolution of PRV in China.<\/jats:p>","DOI":"10.3390\/v15091966","type":"journal-article","created":{"date-parts":[[2023,9,20]],"date-time":"2023-09-20T21:47:03Z","timestamp":1695246423000},"page":"1966","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["Isolation and Characterization of a Novel Recombinant Classical Pseudorabies Virus in the Context of the Variant Strains Pandemic in China"],"prefix":"10.3390","volume":"15","author":[{"given":"Zhengmin","family":"Lian","sequence":"first","affiliation":[{"name":"Jiangsu Co-Innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Panrao","family":"Liu","sequence":"additional","affiliation":[{"name":"Jiangsu Co-Innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenbang","family":"Zhu","sequence":"additional","affiliation":[{"name":"Jiangsu Co-Innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhe","family":"Sun","sequence":"additional","affiliation":[{"name":"Luoyang Putai Biotech Co., Ltd., Luoyang 471003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiuling","family":"Yu","sequence":"additional","affiliation":[{"name":"Luoyang Putai Biotech Co., Ltd., Luoyang 471003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junhua","family":"Deng","sequence":"additional","affiliation":[{"name":"Luoyang Putai Biotech Co., Ltd., Luoyang 471003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruichao","family":"Li","sequence":"additional","affiliation":[{"name":"Jiangsu Co-Innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4635-1602","authenticated-orcid":false,"given":"Xiangdong","family":"Li","sequence":"additional","affiliation":[{"name":"Jiangsu Co-Innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China"},{"name":"Luoyang Putai Biotech Co., Ltd., Luoyang 471003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kegong","family":"Tian","sequence":"additional","affiliation":[{"name":"Luoyang Putai Biotech Co., Ltd., Luoyang 471003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,9,20]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"462","DOI":"10.1128\/MMBR.69.3.462-500.2005","article-title":"Molecular biology of pseudorabies virus: Impact on neurovirology and veterinary medicine","volume":"69","author":"Pomeranz","year":"2005","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1128\/JVI.78.1.424-440.2004","article-title":"Complete, annotated sequence of the pseudorabies virus genome","volume":"78","author":"Klupp","year":"2004","journal-title":"J. Virol."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Szpara, M.L., Tafuri, Y.R., Parsons, L., Shamim, S.R., Verstrepen, K.J., Legendre, M., and Enquist, L.W. (2011). A wide extent of inter-strain diversity in virulent and vaccine strains of alphaherpesviruses. PLoS Pathog., 7.","DOI":"10.1371\/journal.ppat.1002282"},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Zheng, H.H., Fu, P.F., Chen, H.Y., and Wang, Z.Y. (2022). Pseudorabies Virus: From Pathogenesis to Prevention Strategies. Viruses, 14.","DOI":"10.3390\/v14081638"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.1016\/j.vaccine.2017.01.003","article-title":"Bartha-k61 vaccine protects growing pigs against challenge with an emerging variant pseudorabies virus","volume":"35","author":"Zhou","year":"2017","journal-title":"Vaccine"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"102","DOI":"10.3201\/eid2001.130531","article-title":"Pathogenic pseudorabies virus, China, 2012","volume":"20","author":"Yu","year":"2014","journal-title":"Emerg. Infect. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1749","DOI":"10.3201\/eid1911.130177","article-title":"Pseudorabies virus variant in Bartha-K61-vaccinated pigs, China, 2012","volume":"19","author":"An","year":"2013","journal-title":"Emerg. Infect. Dis."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"e5785","DOI":"10.7717\/peerj.5785","article-title":"Epidemiological and genetic characteristics of swine pseudorabies virus in mainland China between 2012 and 2017","volume":"6","author":"Sun","year":"2018","journal-title":"PeerJ"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1454","DOI":"10.1111\/tbed.13813","article-title":"Emergence of a novel pathogenic recombinant virus from Bartha vaccine and variant pseudorabies virus in China","volume":"68","author":"Bo","year":"2021","journal-title":"Transbound. Emerg. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1016\/j.vetmic.2014.09.003","article-title":"Pathogenicity and genomic characterization of a pseudorabies virus variant isolated from Bartha-K61-vaccinated swine population in China","volume":"174","author":"Luo","year":"2014","journal-title":"Vet. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1007\/s12250-020-00340-0","article-title":"Current Status and Challenge of Pseudorabies Virus Infection in China","volume":"36","author":"Tan","year":"2021","journal-title":"Virol. Sin."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1016\/j.vetmic.2015.09.021","article-title":"Emergence of a Pseudorabies virus variant with increased virulence to piglets","volume":"181","author":"Tong","year":"2015","journal-title":"Vet. Microbiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3379","DOI":"10.1016\/j.vaccine.2014.04.035","article-title":"A novel gE-deleted pseudorabies virus (PRV) provides rapid and complete protection from lethal challenge with the PRV variant emerging in Bartha-K61-vaccinated swine population in China","volume":"32","author":"Wang","year":"2014","journal-title":"Vaccine"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"23","DOI":"10.5501\/wjv.v5.i1.23","article-title":"Pathogenicity of a currently circulating Chinese variant pseudorabies virus in pigs","volume":"5","author":"Yang","year":"2016","journal-title":"World J. Virol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.virusres.2014.09.003","article-title":"A novel inactivated gE\/gI deleted pseudorabies virus (PRV) vaccine completely protects pigs from an emerged variant PRV challenge","volume":"195","author":"Gu","year":"2015","journal-title":"Virus Res."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Wang, T., Xiao, Y., Yang, Q., Wang, Y., Sun, Z., Zhang, C., Yan, S., Wang, J., Guo, L., and Yan, H. (2015). Construction of a gE-Deleted Pseudorabies Virus and Its Efficacy to the New-Emerging Variant PRV Challenge in the Form of Killed Vaccine. Biomed. Res. Int., 2015.","DOI":"10.1155\/2015\/684945"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2432","DOI":"10.1016\/j.vaccine.2015.03.094","article-title":"Construction of a triple gene-deleted Chinese Pseudorabies virus variant and its efficacy study as a vaccine candidate on suckling piglets","volume":"33","author":"Zhang","year":"2015","journal-title":"Vaccine"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"797611","DOI":"10.3389\/fvets.2021.797611","article-title":"Construction and Immunogenicity of a Recombinant Pseudorabies Virus Variant With TK\/gI\/gE\/11k\/28k Deletion","volume":"8","author":"Yan","year":"2021","journal-title":"Front. Vet. Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.virol.2016.01.016","article-title":"Genomic analyses reveal that partial sequence of an earlier pseudorabies virus in China is originated from a Bartha-vaccine-like strain","volume":"491","author":"Ye","year":"2016","journal-title":"Virology"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Qin, Y., Qin, S., Huang, X., Xu, L., Ouyang, K., Chen, Y., Wei, Z., and Huang, W. (2023). Isolation and identification of two novel pseudorabies viruses with natural recombination or TK gene deletion in China. Vet. Microbiol., 280.","DOI":"10.1016\/j.vetmic.2023.109703"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"5571","DOI":"10.1128\/JVI.02589-05","article-title":"Influence of pseudorabies virus proteins on neuroinvasion and neurovirulence in mice","volume":"80","author":"Klopfleisch","year":"2006","journal-title":"J. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2956","DOI":"10.1128\/JVI.78.6.2956-2966.2004","article-title":"Influence of tegument proteins of pseudorabies virus on neuroinvasion and transneuronal spread in the nervous system of adult mice after intranasal inoculation","volume":"78","author":"Klopfleisch","year":"2004","journal-title":"J. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.virusres.2017.05.002","article-title":"From mouse to pig: Is PRV vaccine safe across two species?","volume":"236","author":"Li","year":"2017","journal-title":"Virus Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"894","DOI":"10.1128\/jvi.66.2.894-905.1992","article-title":"Pseudorabies virus envelope glycoproteins gp50 and gII are essential for virus penetration, but only gII is involved in membrane fusion","volume":"66","author":"Peeters","year":"1992","journal-title":"J. Virol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"5348","DOI":"10.1128\/jvi.65.10.5348-5356.1991","article-title":"Pseudorabies virus glycoproteins gII and gp50 are essential for virus penetration","volume":"65","author":"Rauh","year":"1991","journal-title":"J. Virol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1099\/0022-1317-80-3-537","article-title":"Distribution of B-cell epitopes on the pseudorabies virus glycoprotein B","volume":"80 Pt 3","author":"Zaripov","year":"1999","journal-title":"J. Gen. Virol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.jviromet.2008.03.015","article-title":"Pseudorabies virus glycoprotein C attachment-proficient revertants isolated through a simple, targeted mutagenesis scheme","volume":"151","author":"Rue","year":"2008","journal-title":"J. Virol. Methods"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Li, A., Lu, G., Qi, J., Wu, L., Tian, K., Luo, T., Shi, Y., Yan, J., and Gao, G.F. (2017). Structural basis of nectin-1 recognition by pseudorabies virus glycoprotein D. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006314"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/j.ijbiomac.2021.07.170","article-title":"A monoclonal antibody neutralizes pesudorabies virus by blocking gD binding to the receptor nectin-1","volume":"188","author":"Zhang","year":"2021","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"10875","DOI":"10.1128\/JVI.79.17.10875-10889.2005","article-title":"Neuron-to-cell spread of pseudorabies virus in a compartmented neuronal culture system","volume":"79","author":"Enquist","year":"2005","journal-title":"J. Virol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"9431","DOI":"10.1128\/JVI.01317-13","article-title":"Glycoproteins gE and gI are required for efficient KIF1A-dependent anterograde axonal transport of alphaherpesvirus particles in neurons","volume":"87","author":"Kratchmarov","year":"2013","journal-title":"J. Virol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Liu, H., Shi, Z., Liu, C., Wang, P., Wang, M., Wang, S., Liu, Z., Wei, L., Sun, Z., and He, X. (2020). Implication of the Identification of an Earlier Pseudorabies Virus (PRV) Strain HLJ-2013 to the Evolution of Chinese PRVs. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.612474"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Huang, J., Tang, W., Wang, X., Zhao, J., Peng, K., Sun, X., Li, S., Kuang, S., Zhu, L., and Zhou, Y. (2022). The Genetic Characterization of a Novel Natural Recombinant Pseudorabies Virus in China. Viruses, 14.","DOI":"10.3390\/v14050978"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Hu, R., Wang, L., Liu, Q., Hua, L., Huang, X., Zhang, Y., Fan, J., Chen, H., Song, W., and Liang, W. (2021). Whole-Genome Sequence Analysis of Pseudorabies Virus Clinical Isolates from Pigs in China between 2012 and 2017 in China. Viruses, 13.","DOI":"10.3390\/v13071322"}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/15\/9\/1966\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T20:54:18Z","timestamp":1760129658000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/15\/9\/1966"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,9,20]]},"references-count":34,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2023,9]]}},"alternative-id":["v15091966"],"URL":"https:\/\/doi.org\/10.3390\/v15091966","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,9,20]]}}}