{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T13:22:02Z","timestamp":1788268922186,"version":"build-2803163510"},"reference-count":54,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T00:00:00Z","timestamp":1787875200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Shaanxi Provincial Natural Science Basic Research Program Key Project","award":["2024JC-ZDXM-42"],"award-info":[{"award-number":["2024JC-ZDXM-42"]}]},{"name":"Shaanxi Provincial Natural Science Basic Research Program Key Project","award":["2025JC-YBMS-988"],"award-info":[{"award-number":["2025JC-YBMS-988"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82272330"],"award-info":[{"award-number":["82272330"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["82272330"],"award-info":[{"award-number":["82272330"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Xijing Hospital","award":["XJZT26JS052"],"award-info":[{"award-number":["XJZT26JS052"]}]},{"award":["XJZT26JS052"],"award-info":[{"award-number":["XJZT26JS052"]}],"id":[{"id":"https:\/\/ror.org\/05cqe9350","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Pathogens"],"abstract":"<jats:p>Herpesviruses establish lifelong persistence through acute replication, immune evasion, latency, and reactivation. Nucleotide-binding oligomerization domain-like receptors (NLRs) are most commonly discussed in the context of inflammasome sensors, pyroptosis, and IL-1 family cytokines. Here, we depart from this canonical framework to focus on three regulatory NLRs\u2014NLRC3, NLRC5, and NLRX1\u2014that act outside inflammasome assembly. Using HSV-1 as the central model and other herpesviruses as comparators, we propose a conceptual framework in which these NLRs modulate three host-control layers: cGAS-STING-dependent DNA sensing, MHC class I antigen presentation, and MAVS-mediated mitochondrial antiviral signaling. We explicitly frame these NLRs as regulatory hypotheses for HSV-1 biology rather than established restriction factors, and we provide prioritized, falsifiable predictions to guide future experimentation. Current evidence is strongest at the pathway level; direct tests of NLRC3, NLRC5, and NLRX1 in acute infection, latency, and reactivation remain necessary. The evidence is asymmetric: NLRC3 has been functionally tested in HSV-1-infected cells and mice, whereas direct HSV-1 regulation by NLRC5 or NLRX1 remains unestablished. Comparative herpesvirus and pathway studies, therefore, support testable hypotheses for NLRC5 and NLRX1 in acute infection, latency and reactivation.<\/jats:p>","DOI":"10.3390\/pathogens15090908","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T12:41:18Z","timestamp":1788266478000},"page":"908","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Regulatory NLRs in HSV-1 Infection: Direct Evidence, Comparative Mechanisms and Testable Hypotheses"],"prefix":"10.3390","volume":"15","author":[{"given":"Mengting","family":"Zhu","sequence":"first","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuai","family":"Zhao","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"},{"name":"School of Medicine, Northwest University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xing","family":"Sun","sequence":"additional","affiliation":[{"name":"School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuhan","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shiyuan","family":"Hou","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xing","family":"Shen","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jielin","family":"Zhou","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ziyu","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xingan","family":"Wu","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9725-3463","authenticated-orcid":false,"given":"Rongrong","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Microbiology, School of Basic Medical Sciences, Fourth Military Medical University, Xi\u2019an 710032, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2425744","DOI":"10.1080\/21505594.2024.2425744","article-title":"An updated review of HSV-1 infection-associated diseases and treatment, vaccine development, and vector therapy application","volume":"15","author":"Su","year":"2024","journal-title":"Virulence"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Zhao, J., Qin, C., Liu, Y., Rao, Y., and Feng, P. (2020). Herpes Simplex Virus and Pattern Recognition Receptors: An Arms Race. Front. Immunol., 11.","DOI":"10.3389\/fimmu.2020.613799"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1016\/j.immuni.2014.01.010","article-title":"NLRC3, a member of the NLR family of proteins, is a negative regulator of innate immune signaling induced by the DNA sensor STING","volume":"40","author":"Zhang","year":"2014","journal-title":"Immunity"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"13794","DOI":"10.1073\/pnas.1008684107","article-title":"NLR family member NLRC5 is a transcriptional regulator of MHC class I genes","volume":"107","author":"Meissner","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1038\/nri3339","article-title":"NLRC5: A key regulator of MHC class I-dependent immune responses","volume":"12","author":"Kobayashi","year":"2012","journal-title":"Nat. Rev. Immunol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1038\/nature06501","article-title":"NLRX1 is a regulator of mitochondrial antiviral immunity","volume":"451","author":"Moore","year":"2008","journal-title":"Nature"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1016\/j.immuni.2012.03.025","article-title":"The mitochondrial proteins NLRX1 and TUFM form a complex that regulates type I interferon and autophagy","volume":"36","author":"Lei","year":"2012","journal-title":"Immunity"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1038\/s41392-025-02264-1","article-title":"Pattern recognition receptors: Function, regulation and therapeutic potential","volume":"10","author":"Chen","year":"2025","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1038\/s41577-023-00849-x","article-title":"The NLR gene family: From discovery to present day","volume":"23","author":"Chou","year":"2023","journal-title":"Nat. Rev. Immunol."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Kienes, I., Weidl, T., Mirza, N., Chamaillard, M., and Kufer, T.A. (2021). Role of NLRs in the Regulation of Type I Interferon Signaling, Host Defense and Tolerance to Inflammation. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22031301"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"e13429","DOI":"10.1111\/imr.13429","article-title":"Innate Immunity Never \u201cNODs\u201d Off: NLRs Regulate the Host Anti-Viral Immune Response","volume":"330","author":"Woolls","year":"2025","journal-title":"Immunol. Rev."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Wang, W., Hu, D., Wu, C., Feng, Y., Li, A., Liu, W., Wang, Y., Chen, K., Tian, M., and Xiao, F. (2020). STING promotes NLRP3 localization in ER and facilitates NLRP3 deubiquitination to activate the inflammasome upon HSV-1 infection. PLoS Pathog., 16.","DOI":"10.1371\/journal.ppat.1008335"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"602","DOI":"10.1016\/j.chom.2019.02.013","article-title":"NLRP12 Regulates Anti-viral RIG-I Activation via Interaction with TRIM25","volume":"25","author":"Chen","year":"2019","journal-title":"Cell Host Microbe"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Jiang, J., Zhang, D., Liu, W., Yang, J., Yang, F., Liu, J., and Hu, K. (2024). Overexpression of NLRP12 enhances macrophage immune response and alleviates herpes simplex keratitis. Front. Cell. Infect. Microbiol., 14.","DOI":"10.3389\/fcimb.2024.1416105"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Fekete, T., Bencze, D., Szabo, A., Csoma, E., Biro, T., Bacsi, A., and Pazmandi, K. (2018). Regulatory NLRs Control the RLR-Mediated Type I Interferon and Inflammatory Responses in Human Dendritic Cells. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.02314"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"4418","DOI":"10.1038\/s41467-025-59338-5","article-title":"TRIM23 mediates cGAS-induced autophagy in anti-HSV defense","volume":"16","author":"Acharya","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"102051","DOI":"10.1016\/j.xcrm.2025.102051","article-title":"Herpes simplex virus 1 encodes a STING antagonist that can be therapeutically targeted","volume":"6","author":"Wang","year":"2025","journal-title":"Cell Rep. Med."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"763","DOI":"10.1080\/15548627.2026.2614901","article-title":"Alphaherpesvirus UL48 homologs degrade STING1 by selective autophagy","volume":"22","author":"Kong","year":"2026","journal-title":"Autophagy"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Qu, W., Yuan, Y., Shen, D., Zhang, J., Huang, X., Wu, L., Yin, H., Zhao, Z., Wang, H., and Chai, L. (2025). Alpha-herpesvirus US1 interacts with cGAS to suppress type I IFN responses and antiviral defense. PLoS Pathog., 21.","DOI":"10.1371\/journal.ppat.1013669"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"110604","DOI":"10.1016\/j.vetmic.2025.110604","article-title":"Pseudorabies virus DNA binding to porcine NLRC3 unleashes the activation of the porcine NLRP3 inflammasome","volume":"307","author":"Li","year":"2025","journal-title":"Vet. Microbiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"e29504","DOI":"10.1002\/jmv.29504","article-title":"Dynamic bidirectional regulation of NLRC3 and gammaherpesviruses during viral latency in B lymphocytes","volume":"96","author":"Kang","year":"2024","journal-title":"J. Med. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Sun, D., Xu, J., Zhang, W., Song, C., Gao, C., He, Y., and Shang, Y. (2022). Negative regulator NLRC3: Its potential role and regulatory mechanism in immune response and immune-related diseases. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.1012459"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"111145","DOI":"10.1016\/j.isci.2024.111145","article-title":"Porcine NLRC3 specially binds short dsDNA to regulate cGAS activation","volume":"27","author":"Li","year":"2024","journal-title":"iScience"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e0388125","DOI":"10.1128\/mbio.03881-25","article-title":"Linker histone H1.2 inhibits HSV-1-induced IFN response via cGAS","volume":"17","author":"Li","year":"2026","journal-title":"mBio"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1186\/s12974-026-03856-6","article-title":"LAPF enhances lysosomal acidification to promote TLR9 and cGAS-STING-mediated antiviral immunity and attenuate HSV-1-induced neuroinflammatory pain","volume":"23","author":"Kong","year":"2026","journal-title":"J. Neuroinflammation"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1093\/infdis\/jiaf404","article-title":"Histone Deacetylase 6 Regulates \u03b1-Tubulin to Inhibit the Stimulator of Interferon Genes\/NLR Family Pyrin Domain-Containing 3-Mediated Microglial Pyroptosis Induced by Herpes Simplex Virus Type 1 Infection","volume":"233","author":"Zeng","year":"2026","journal-title":"J. Infect. Dis."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"110351","DOI":"10.1016\/j.fsi.2025.110351","article-title":"NLRC3 regulates RIP2, STING, TBK1, and TRAF6 mediated type I IFN signaling and inflammatory response in large yellow croaker Larimichthys crocea","volume":"162","author":"Xu","year":"2025","journal-title":"Fish. Shellfish Immunol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"822","DOI":"10.1038\/cr.2012.53","article-title":"Enhanced TLR-induced NF-\u03baB signaling and type I interferon responses in NLRC5 deficient mice","volume":"22","author":"Tong","year":"2012","journal-title":"Cell Res."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Godkowicz, M., and Druszczy\u0144ska, M. (2022). NOD1, NOD2, and NLRC5 Receptors in Antiviral and Antimycobacterial Immunity. Vaccines, 10.","DOI":"10.3390\/vaccines10091487"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1186\/s12964-024-02012-y","article-title":"New insights into the structure domain and function of NLR family CARD domain containing 5","volume":"23","author":"Zhu","year":"2025","journal-title":"Cell Commun. Signal."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Brunschwiler, F., Nakka, S., Guerra, J., and Guarda, G. (2024). A M\u00e9nage \u00e0 trois: NLRC5, immunity, and metabolism. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1426620"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"107205","DOI":"10.1016\/j.jbc.2024.107205","article-title":"The balance between nuclear import and export of NLRC5 regulates MHC class I transactivation","volume":"300","author":"Zhu","year":"2024","journal-title":"J. Biol. Chem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e2300109","DOI":"10.1002\/bies.202300109","article-title":"NLRC5\/MHC class I transactivator: A key target for immune escape by SARS-CoV-2","volume":"46","author":"Zhu","year":"2024","journal-title":"Bioessays"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"6602","DOI":"10.1038\/s41467-021-26910-8","article-title":"SARS-CoV-2 inhibits induction of the MHC class I pathway by targeting the STAT1-IRF1-NLRC5 axis","volume":"12","author":"Yoo","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"108801","DOI":"10.1016\/j.isci.2024.108801","article-title":"Multi-targeted loss of the antigen presentation molecule MR1 during HSV-1 and HSV-2 infection","volume":"27","author":"Samer","year":"2024","journal-title":"iScience"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"107748","DOI":"10.1016\/j.jbc.2024.107748","article-title":"Impaired endocytosis and accumulation in early endosomal compartments defines herpes simplex virus-mediated disruption of the nonclassical MHC class I-related molecule MR1","volume":"300","author":"Samer","year":"2024","journal-title":"J. Biol. Chem."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Croft, N.P., Shannon-Lowe, C., Bell, A.I., Horst, D., Kremmer, E., Ressing, M.E., Wiertz, E.J., Middeldorp, J.M., Rowe, M., and Rickinson, A.B. (2009). Stage-specific inhibition of MHC class I presentation by the Epstein-Barr virus BNLF2a protein during virus lytic cycle. PLoS Pathog., 5.","DOI":"10.1371\/journal.ppat.1000490"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1128\/JVI.02183-15","article-title":"The Missing Link in Epstein-Barr Virus Immune Evasion: The BDLF3 Gene Induces Ubiquitination and Downregulation of Major Histocompatibility Complex Class I (MHC-I) and MHC-II","volume":"90","author":"Quinn","year":"2016","journal-title":"J. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Apcher, S., Daskalogianni, C., Manoury, B., and F\u00e5hraeus, R. (2010). Epstein Barr virus-encoded EBNA1 interference with MHC class I antigen presentation reveals a close correlation between mRNA translation initiation and antigen presentation. PLoS Pathog., 6.","DOI":"10.1371\/journal.ppat.1001151"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Zuo, J., Currin, A., Griffin, B.D., Shannon-Lowe, C., Thomas, W.A., Ressing, M.E., Wiertz, E.J., and Rowe, M. (2009). The Epstein-Barr virus G-protein-coupled receptor contributes to immune evasion by targeting MHC class I molecules for degradation. PLoS Pathog., 5.","DOI":"10.1371\/journal.ppat.1000255"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"8051","DOI":"10.1073\/pnas.140129797","article-title":"Kaposi\u2019s sarcoma-associated herpesvirus encodes two proteins that block cell surface display of MHC class I chains by enhancing their endocytosis","volume":"97","author":"Coscoy","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"5300","DOI":"10.1128\/JVI.74.11.5300-5309.2000","article-title":"Downregulation of major histocompatibility complex class I molecules by Kaposi\u2019s sarcoma-associated herpesvirus K3 and K5 proteins","volume":"74","author":"Ishido","year":"2000","journal-title":"J. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Gabor, F., Jahn, G., Sedmak, D.D., and Sinzger, C. (2020). In vivo Downregulation of MHC Class I Molecules by HCMV Occurs During All Phases of Viral Replication but Is Not Always Complete. Front. Cell. Infect. Microbiol., 10.","DOI":"10.3389\/fcimb.2020.00283"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2061","DOI":"10.1038\/s41467-021-21983-x","article-title":"A slowly cleaved viral signal peptide acts as a protein-integral immune evasion domain","volume":"12","author":"Seidel","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Nagai-Singer, M.A., Morrison, H.A., and Allen, I.C. (2019). NLRX1 Is a Multifaceted and Enigmatic Regulator of Immune System Function. Front. Immunol., 10.","DOI":"10.3389\/fimmu.2019.02419"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e01264-17","DOI":"10.1128\/JVI.01264-17","article-title":"NLRX1 Mediates MAVS Degradation To Attenuate the Hepatitis C Virus-Induced Innate Immune Response through PCBP2","volume":"91","author":"Qin","year":"2017","journal-title":"J. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Liu, X., Ma, Y., Voss, K., van Gent, M., Chan, Y.K., Gack, M.U., Gale, M., and He, B. (2021). The herpesvirus accessory protein \u03b3134.5 facilitates viral replication by disabling mitochondrial translocation of RIG-I. PLoS Pathog., 17.","DOI":"10.1371\/journal.ppat.1009446"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"eabd0276","DOI":"10.1126\/sciadv.abd0276","article-title":"PPM1G restricts innate immune signaling mediated by STING and MAVS and is hijacked by KSHV for immune evasion","volume":"6","author":"Yu","year":"2020","journal-title":"Sci. Adv."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1041","DOI":"10.1038\/s41564-022-01136-6","article-title":"Dual inhibition of innate immunity and apoptosis by human cytomegalovirus protein UL37x1 enables efficient virus replication","volume":"7","author":"Ren","year":"2022","journal-title":"Nat. Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Woolls, M.K., Elliott, C.M., Ivester, H.M., and Allen, I.C. (2025). Mystery machine: The complex roles of NLRX1 in viral infection. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1581313"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.tim.2022.07.008","article-title":"T cell immunity in HSV-1- and VZV-infected neural ganglia","volume":"31","author":"Boeren","year":"2023","journal-title":"Trends Microbiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"115741","DOI":"10.1016\/j.celrep.2025.115741","article-title":"KSHV miRNAs target STING to evade innate immunity and facilitate KSHV lytic reactivation from latency","volume":"44","author":"Paulsen","year":"2025","journal-title":"Cell Rep."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"9015","DOI":"10.1038\/s41467-025-64037-2","article-title":"Extracellular vesicles derived EBV tegument protein BRRF2 suppresses cGAS phase separation to promote anti-viral innate immune evasion","volume":"16","author":"Hu","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/j.chom.2015.07.015","article-title":"Inhibition of cGAS DNA Sensing by a Herpesvirus Virion Protein","volume":"18","author":"Wu","year":"2015","journal-title":"Cell Host Microbe"}],"container-title":["Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/9\/908\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T12:57:49Z","timestamp":1788267469000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/9\/908"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,28]]},"references-count":54,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["pathogens15090908"],"URL":"https:\/\/doi.org\/10.3390\/pathogens15090908","relation":{},"ISSN":["2076-0817"],"issn-type":[{"value":"2076-0817","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,28]]}}}