{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T11:15:48Z","timestamp":1788434148620,"version":"build-2803163510"},"reference-count":75,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T00:00:00Z","timestamp":1788393600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Vaccines"],"abstract":"<jats:p>Background\/Objectives: Live attenuated mumps virus (MuV) vaccine strains have significantly reduced disease incidence since their introduction in the 1960s; however, a recent resurgence of outbreaks has led to calls for the development of new vaccines to overcome what appears to be reduced effectiveness linked to waning immunity and the emergence of strains antigenically mismatched to vaccine strains. A major obstacle to the development of newer live, attenuated MuV vaccines is ensuring their safety, particularly given the virus\u2019s neurotropic properties. Indeed, several vaccine strains licensed for use outside the US have proven to be insufficiently attenuated (such as the Urabe AM9 vaccine strain) and have caused aseptic meningitis in recipients, despite efforts to test these strains for neurotoxicity pre-licensure. Historically, attenuation was achieved empirically, and pre-clinical testing for neurovirulence safety has proven unreliable. Despite efforts in recent years, the genetic basis of attenuation of MuV vaccines remains inadequately understood. The objective of this study was to elucidate the genetic basis of neurovirulence of the Urabe AM9 vaccine strain. Methods: To this end, we generated a series of chimeric viruses in which genes of the highly attenuated Jeryl Lynn (JL) vaccine strain were exchanged with corresponding genes from the Urabe AM9 vaccine strain. The resulting chimeric viruses were tested for neurovirulence in a rat model and characterized for replication in vitro and in vivo. Using a multi-tiered approach consisting of independent rescue and analysis of two to three viruses per cDNA construct, possible off-target effects of identified single-nucleotide heterogeneities were mitigated. Results: All viruses were shown to be replication-competent in Vero cells, but differences in replication efficiencies and virus-induced neurotoxicity were observed in the in vivo model. In a rat neuronal cell line, the Urabe AM9 M and HN genes had opposite effects on the growth of chimeric JL- and Urabe AM9-based viruses. Conclusions: The results presented herein indicate that MuV neuroattenuation is mediated by the concerted action of multiple genes, with the matrix (M) gene exerting a dominant effect. These findings contribute to a growing body of evidence that informs the rational design of next-generation live-attenuated mumps virus vaccines.<\/jats:p>","DOI":"10.3390\/vaccines14090774","type":"journal-article","created":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T10:10:33Z","timestamp":1788430233000},"page":"774","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A Chimeric Virus Approach Reveals the Matrix (M) Gene as a Critical Modulator of Mumps Virus Neurovirulence"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5490-402X","authenticated-orcid":false,"given":"Christian J.","family":"Sauder","sequence":"first","affiliation":[{"name":"Division of Viral Products, Office of Vaccine Research and Review, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"given":"Malen","family":"Link","sequence":"additional","affiliation":[{"name":"Division of Viral Products, Office of Vaccine Research and Review, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, 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20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Chao-Kai","family":"Chou","sequence":"additional","affiliation":[{"name":"Facility for Biotechnology Resources, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6392-2235","authenticated-orcid":false,"given":"Wells W.","family":"Wu","sequence":"additional","affiliation":[{"name":"Facility for Biotechnology Resources, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Tatiana","family":"Zagorodnyaya","sequence":"additional","affiliation":[{"name":"Division of Viral Products, Office of Vaccine Research and Review, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0110-5504","authenticated-orcid":false,"given":"Majid","family":"Laassri","sequence":"additional","affiliation":[{"name":"Division of Viral Products, Office of Vaccine Research and Review, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Steven","family":"Rubin","sequence":"additional","affiliation":[{"name":"Division of Viral Products, Office of Vaccine Research and Review, Center for Biologics Evaluation and Research (CBER), Food and Drug Administration (FDA), Silver Spring, MD 20993, USA"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"3110","DOI":"10.1080\/21645515.2016.1210745","article-title":"Mumps virus pathogenesis: Insights and knowledge gaps","volume":"12","author":"Gouma","year":"2016","journal-title":"Hum. Vaccines Immunother."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Su, S.B., Chang, H.L., and Chen, A.K. (2020). Current Status of Mumps Virus Infection: Epidemiology, Pathogenesis, and Vaccine. Int. J. Environ. Res. Public Health, 17.","DOI":"10.3390\/ijerph17051686"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"eaao5945","DOI":"10.1126\/scitranslmed.aao5945","article-title":"Vaccine waning and mumps re-emergence in the United States","volume":"10","author":"Lewnard","year":"2018","journal-title":"Sci. Transl. Med."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Connell, A.R., Connell, J., Leahy, T.R., and Hassan, J. (2020). Mumps Outbreaks in Vaccinated Populations-Is It Time to Re-assess the Clinical Efficacy of Vaccines?. Front. Immunol., 11.","DOI":"10.3389\/fimmu.2020.02089"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"126143","DOI":"10.1016\/j.vaccine.2024.07.044","article-title":"Characteristics of reported mumps cases in the United States: 2018-2023","volume":"42","author":"Tappe","year":"2024","journal-title":"Vaccine"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"e2207595120","DOI":"10.1073\/pnas.2207595120","article-title":"Disentangling the causes of mumps reemergence in the United States","volume":"120","author":"Gokhale","year":"2023","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"19071","DOI":"10.1073\/pnas.1905570116","article-title":"Decreased humoral immunity to mumps in young adults immunized with MMR vaccine in childhood","volume":"116","author":"Rasheed","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"13337","DOI":"10.1038\/s41598-018-31630-z","article-title":"Differences in antigenic sites and other functional regions between genotype A and G mumps virus surface proteins","volume":"8","author":"Gouma","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"3988","DOI":"10.1016\/j.vaccine.2017.05.084","article-title":"Immunogenicity of mumps virus vaccine candidates matching circulating genotypes in the United States and China","volume":"35","author":"Zengel","year":"2017","journal-title":"Vaccine"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"ofae329","DOI":"10.1093\/ofid\/ofae329","article-title":"Identifying a Level of Neutralizing Antibody That Correlates With Protection From Clinical Mumps Disease During a 2017 Mumps Outbreak Among Military Service Members","volume":"11","author":"Sowers","year":"2024","journal-title":"Open Forum Infect. Dis."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1097\/INF.0000000000004241","article-title":"We Need a New Mumps Vaccine","volume":"43","author":"Plotkin","year":"2024","journal-title":"Pediatr. Infect. Dis. J."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/j.japh.2018.07.004","article-title":"Mumps outbreaks: Implementation of a third dose of mumps-containing vaccine","volume":"58","author":"Berlin","year":"2018","journal-title":"J. Am. Pharm. Assoc."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e00151-19","DOI":"10.1128\/CMR.00151-19","article-title":"Mumps: An Update on Outbreaks, Vaccine Efficacy, and Genomic Diversity","volume":"33","author":"Lam","year":"2020","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"e0198321","DOI":"10.1128\/jvi.01983-21","article-title":"Immunogenicity of Mumps Virus Genotype G Vaccine Candidates in Jeryl Lynn-Immunized Mice","volume":"96","author":"Briggs","year":"2022","journal-title":"J. Virol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Kim, S.Y., Won, H., Hwang, Y.H., Kim, S.E., Lee, J.A., Kim, D., Kim, Y.J., and Lee, T.Y. (2024). Immunogenicity and Cross-Protection Efficacy of a Genotype F-Derived Attenuated Virus Vaccine Candidate against Mumps Virus in Mice. Vaccines, 12.","DOI":"10.3390\/vaccines12060595"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"e2404053121","DOI":"10.1073\/pnas.2404053121","article-title":"Structure-based design of glycoprotein subunit vaccines for mumps","volume":"121","author":"Loomis","year":"2024","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2698180","DOI":"10.1080\/21645515.2026.2698180","article-title":"Mumps resurgence despite MMR: Limitations of current vaccines and the need for new strategies","volume":"22","author":"Vashishtha","year":"2026","journal-title":"Hum. Vaccines Immunother."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"17264","DOI":"10.1073\/pnas.0407639101","article-title":"The V proteins of paramyxoviruses bind the IFN-inducible RNA helicase, mda-5, and inhibit its activation of the IFN-beta promoter","volume":"101","author":"Andrejeva","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"12676","DOI":"10.1128\/JVI.76.24.12676-12682.2002","article-title":"Association of mumps virus V protein with RACK1 results in dissociation of STAT-1 from the alpha interferon receptor complex","volume":"76","author":"Kubota","year":"2002","journal-title":"J. Virol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1007\/s00430-017-0509-y","article-title":"Tetherin antagonism by V proteins is a common trait among the genus Rubulavirus","volume":"206","author":"Ohta","year":"2017","journal-title":"Med. Microbiol. Immunol."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Risalvato, J. (2026). Mumps Virus: Replication, Immune Response, and the Changing Landscape of Vaccine Effectiveness. Pathogens, 15.","DOI":"10.3390\/pathogens15010072"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1416","DOI":"10.1016\/j.biocel.2010.04.005","article-title":"Paramyxovirus assembly and budding: Building particles that transmit infections","volume":"42","author":"Harrison","year":"2010","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"7261","DOI":"10.1128\/JVI.00421-09","article-title":"Mumps virus matrix, fusion, and nucleocapsid proteins cooperate for efficient production of virus-like particles","volume":"83","author":"Li","year":"2009","journal-title":"J. Virol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"eads3071","DOI":"10.1126\/sciadv.ads3071","article-title":"The SH protein of mumps virus is a druggable pentameric viroporin","volume":"11","author":"Devantier","year":"2025","journal-title":"Sci. Adv."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"e01037-17","DOI":"10.1128\/JVI.01037-17","article-title":"Mumps Virus SH Protein Inhibits NF-\u03baB Activation by Interacting with Tumor Necrosis Factor Receptor 1, Interleukin-1 Receptor 1, and Toll-Like Receptor 3 Complexes","volume":"91","author":"Franz","year":"2017","journal-title":"J. Virol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.virol.2011.05.003","article-title":"Rescue of wild-type mumps virus from a strain associated with recent outbreaks helps to define the role of the SH ORF in the pathogenesis of mumps virus","volume":"417","author":"Xu","year":"2011","journal-title":"Virology"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1006\/viro.1996.0583","article-title":"The mumps virus SH protein is a membrane protein and not essential for virus growth","volume":"225","author":"Takeuchi","year":"1996","journal-title":"Virology"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"6082","DOI":"10.1128\/JVI.02686-10","article-title":"Discrimination of mumps virus small hydrophobic gene deletion effects from gene translation effects on virus virulence","volume":"85","author":"Malik","year":"2011","journal-title":"J. Virol."},{"key":"ref_29","unstructured":"Knipe, D.M., and Howley, P. (2013). Paramyxoviridae: The viruses and their replication. Fields Virology, Williams and Wilkins. [6th ed.]."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1086\/314905","article-title":"The mumps virus neurovirulence safety test in Rhesus monkeys: A comparison of mumps virus strains","volume":"180","author":"Rubin","year":"1999","journal-title":"J. Infect. Dis."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1123","DOI":"10.1086\/428098","article-title":"The rat-based neurovirulence safety test for the assessment of mumps virus neurovirulence in humans: An international collaborative study","volume":"191","author":"Rubin","year":"2005","journal-title":"J. Infect. Dis."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1111\/j.1442-200X.1996.tb03471.x","article-title":"Adverse events associated with MMR vaccines in Japan","volume":"38","author":"Kimura","year":"1996","journal-title":"Acta Paediatr. Jpn."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"7037","DOI":"10.1016\/j.vaccine.2006.06.049","article-title":"Mumps vaccine virus strains and aseptic meningitis","volume":"24","author":"Bonnet","year":"2006","journal-title":"Vaccine"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1002\/(SICI)1099-1654(199807\/09)8:3<129::AID-RMV213>3.0.CO;2-Z","article-title":"Genetic studies on a mumps vaccine strain associated with meningitis","volume":"8","author":"Brown","year":"1998","journal-title":"Rev. Med. Virol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1093\/oxfordjournals.aje.a010239","article-title":"Outbreak of aseptic meningitis associated with mass vaccination with a urabe-containing measles-mumps-rubella vaccine: Implications for immunization programs","volume":"151","author":"Dourado","year":"2000","journal-title":"Am. J. Epidemiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2850","DOI":"10.1016\/j.vaccine.2011.02.005","article-title":"Neurovirulence safety testing of mumps vaccines--historical perspective and current status","volume":"29","author":"Rubin","year":"2011","journal-title":"Vaccine"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"5382","DOI":"10.1128\/JVI.74.11.5382-5384.2000","article-title":"Evaluation of a neonatal rat model for prediction of mumps virus neurovirulence in humans","volume":"74","author":"Rubin","year":"2000","journal-title":"J. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"5822","DOI":"10.1016\/j.vaccine.2009.07.051","article-title":"Presence of lysine at aa 335 of the hemagglutinin-neuraminidase protein of mumps virus vaccine strain Urabe AM9 is not a requirement for neurovirulence","volume":"27","author":"Sauder","year":"2009","journal-title":"Vaccine"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"7059","DOI":"10.1128\/JVI.00245-11","article-title":"Gene-specific contributions to mumps virus neurovirulence and neuroattenuation","volume":"85","author":"Sauder","year":"2011","journal-title":"J. Virol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2383","DOI":"10.1016\/j.vaccine.2016.03.033","article-title":"Influence of population diversity on neurovirulence potential of plaque purified L-Zagreb variants","volume":"34","author":"Forcic","year":"2016","journal-title":"Vaccine"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"8293","DOI":"10.1128\/JVI.00266-07","article-title":"The F gene of rodent brain-adapted mumps virus is a major determinant of neurovirulence","volume":"81","author":"Lemon","year":"2007","journal-title":"J. Virol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1002\/jmv.21381","article-title":"Identification of genetic mutations associated with attenuation and changes in tropism of Urabe mumps virus","volume":"81","author":"Shah","year":"2009","journal-title":"J. Med. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1080\/13550280701658382","article-title":"A single nucleotide change in the mumps virus F gene affects virus fusogenicity in vitro and virulence in vivo","volume":"13","author":"Malik","year":"2007","journal-title":"J. NeuroVirology"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1768","DOI":"10.1128\/JVI.06019-11","article-title":"The V protein of mumps virus plays a critical role in pathogenesis","volume":"86","author":"Xu","year":"2012","journal-title":"J. Virol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2600","DOI":"10.1128\/JVI.02778-13","article-title":"Immunogenicity of novel mumps vaccine candidates generated by genetic modification","volume":"88","author":"Xu","year":"2014","journal-title":"J. Virol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1002\/rmv.1819","article-title":"Genomic diversity of mumps virus and global distribution of the 12 genotypes","volume":"25","author":"Jin","year":"2015","journal-title":"Rev. Med. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1006\/viro.2001.1009","article-title":"Correlation of genetic variability with safety of mumps vaccine Urabe AM9 strain","volume":"287","author":"Amexis","year":"2001","journal-title":"Virology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3743","DOI":"10.1128\/JCM.36.12.3743-3744.1998","article-title":"Nucleotide sequence at position 1081 of the hemagglutinin-neuraminidasegene in wild-type strains of mumps virus is the most relevant marker of virulence","volume":"36","author":"Cusi","year":"1998","journal-title":"J. Clin. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1093\/infdis\/174.3.619","article-title":"The Urabe AM9 mumps vaccine is a mixture of viruses differing at amino acid 335 of the hemagglutinin-neuraminidase gene with one form associated with disease","volume":"174","author":"Brown","year":"1996","journal-title":"J. Infect. Dis."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/S0168-1702(00)00129-5","article-title":"Biological characteristics of genetic variants of Urabe AM9 mumps vaccine virus","volume":"67","author":"Wright","year":"2000","journal-title":"Virus Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.virol.2006.01.035","article-title":"Changes in mumps virus neurovirulence phenotype associated with quasispecies heterogeneity","volume":"350","author":"Sauder","year":"2006","journal-title":"Virology"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"4831","DOI":"10.1128\/jvi.74.10.4831-4838.2000","article-title":"Rescue of mumps virus from cDNA","volume":"74","author":"Clarke","year":"2000","journal-title":"J. Virol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1128\/JVI.73.1.251-259.1999","article-title":"Generation of bovine respiratory syncytial virus (BRSV) from cDNA: BRSV NS2 is not essential for virus replication in tissue culture, and the human RSV leader region acts as a functional BRSV genome promoter","volume":"73","author":"Buchholz","year":"1999","journal-title":"J. Virol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1006\/exnr.2002.7898","article-title":"AF5, a CNS cell line immortalized with an N-terminal fragment of SV40 large T: Growth, differentiation, genetic stability, and gene expression","volume":"175","author":"Truckenmiller","year":"2002","journal-title":"Exp. Neurol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/BF02789105","article-title":"PCR-mediated recombination and mutagenesis. SOEing together tailor-made genes","volume":"3","author":"Horton","year":"1995","journal-title":"Mol. Biotechnol."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Santana-Quintero, L., Dingerdissen, H., Thierry-Mieg, J., Mazumder, R., and Simonyan, V. (2014). HIVE-hexagon: High-performance, parallelized sequence alignment for next-generation sequencing data analysis. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0099033"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"957","DOI":"10.3390\/genes5040957","article-title":"High-Performance Integrated Virtual Environment (HIVE) Tools and Applications for Big Data Analysis","volume":"5","author":"Simonyan","year":"2014","journal-title":"Genes"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Slovi\u0107, A., Ko\u0161uti\u0107-Gulija, T., For\u010di\u0107, D., \u0160antak, M., Jagu\u0161i\u0107, M., Jurkovi\u0107, M., Pali, D., and Ivan\u010di\u0107-Jele\u010dki, J. (2021). Population Variability Generated during Rescue Process and Passaging of Recombinant Mumps Viruses. Viruses, 13.","DOI":"10.3390\/v13122550"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1099\/vir.0.009449-0","article-title":"Single amino acid changes in the mumps virus haemagglutinin-neuraminidase and polymerase proteins are associated with neuroattenuation","volume":"90","author":"Malik","year":"2009","journal-title":"J. Gen. Virol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1007\/s11262-015-1204-y","article-title":"Generation and propagation of recombinant mumps viruses exhibiting an additional U residue in the homopolymeric U tract of the F gene-end signal","volume":"51","author":"Sauder","year":"2015","journal-title":"Virus Genes"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1016\/j.virol.2016.03.021","article-title":"Evidence that a polyhexameric genome length is preferred, but not strictly required, for efficient mumps virus replication","volume":"493","author":"Sauder","year":"2016","journal-title":"Virology"},{"key":"ref_62","first-page":"CD004407","article-title":"Vaccines for measles, mumps, rubella, and varicella in children","volume":"11","author":"Rivetti","year":"2021","journal-title":"Cochrane Database Syst. Rev."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1186\/s12985-015-0366-5","article-title":"The M, F and HN genes of genotype VIId Newcastle disease virus are associated with the severe pathological changes in the spleen of chickens","volume":"12","author":"Kai","year":"2015","journal-title":"Virol. J."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"7162","DOI":"10.1128\/JVI.00051-11","article-title":"Canine distemper virus matrix protein influences particle infectivity, particle composition, and envelope distribution in polarized epithelial cells and modulates virulence","volume":"85","author":"Dietzel","year":"2011","journal-title":"J. Virol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.virusres.2004.08.010","article-title":"Molecular mechanism of paramyxovirus budding","volume":"106","author":"Takimoto","year":"2004","journal-title":"Virus Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"e0411622","DOI":"10.1128\/spectrum.04116-22","article-title":"Mutation of Phenylalanine 23 of Newcastle Disease Virus Matrix Protein Inhibits Virus Release by Disrupting the Interaction between the FPIV L-Domain and Charged Multivesicular Body Protein 4B","volume":"11","author":"Pei","year":"2023","journal-title":"Microbiol. Spectr."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Ray, G., Schmitt, P.T., and Schmitt, A.P. (2019). Angiomotin-Like 1 Links Paramyxovirus M Proteins to NEDD4 Family Ubiquitin Ligases. Viruses, 11.","DOI":"10.3390\/v11020128"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Pentecost, M., Vashisht, A.A., Lester, T., Voros, T., Beaty, S.M., Park, A., Wang, Y.E., Yun, T.E., Freiberg, A.N., and Wohlschlegel, J.A. (2015). Evidence for ubiquitin-regulated nuclear and subnuclear trafficking among Paramyxovirinae matrix proteins. PLoS Pathog., 11.","DOI":"10.1371\/journal.ppat.1004739"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"e0072222","DOI":"10.1128\/jvi.00722-22","article-title":"Nucleolar Protein Treacle Is Important for the Efficient Growth of Mumps Virus","volume":"96","author":"Wakata","year":"2022","journal-title":"J. Virol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"3650","DOI":"10.1128\/JVI.02673-15","article-title":"C-Terminal DxD-Containing Sequences within Paramyxovirus Nucleocapsid Proteins Determine Matrix Protein Compatibility and Can Direct Foreign Proteins into Budding Particles","volume":"90","author":"Ray","year":"2016","journal-title":"J. Virol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"11616","DOI":"10.1128\/JVI.77.21.11616-11624.2003","article-title":"Changes in mumps virus gene sequence associated with variability in neurovirulent phenotype","volume":"77","author":"Rubin","year":"2003","journal-title":"J. Virol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"6160","DOI":"10.1016\/j.vaccine.2009.08.020","article-title":"Amino acid substitution at position 464 in the haemagglutinin-neuraminidase protein of a mumps virus Urabe strain enhanced the virus growth in neuroblastoma SH-SY5Y cells","volume":"27","author":"Ninomiya","year":"2009","journal-title":"Vaccine"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1006\/viro.1993.1102","article-title":"Canine distemper virus L gene: Sequence and comparison with related viruses","volume":"193","author":"Sidhu","year":"1993","journal-title":"Virology"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1099\/0022-1317-71-5-1153","article-title":"Sequence comparison of five polymerases (L proteins) of unsegmented negative-strand RNA viruses: Theoretical assignment of functional domains","volume":"71","author":"Poch","year":"1990","journal-title":"J. Gen. Virol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"126487","DOI":"10.1016\/j.vaccine.2024.126487","article-title":"Mumps virus neurovirulence assessment-impact of viral doses, animal sex and age on results dispersion","volume":"43","author":"Balija","year":"2025","journal-title":"Vaccine"}],"container-title":["Vaccines"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-393X\/14\/9\/774\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T10:28:16Z","timestamp":1788431296000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-393X\/14\/9\/774"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,3]]},"references-count":75,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["vaccines14090774"],"URL":"https:\/\/doi.org\/10.3390\/vaccines14090774","relation":{},"ISSN":["2076-393X"],"issn-type":[{"value":"2076-393X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,3]]}}}