{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,18]],"date-time":"2026-07-18T17:17:43Z","timestamp":1784395063188,"version":"3.55.0"},"reference-count":44,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T00:00:00Z","timestamp":1777593600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/501100006407","name":"Henan Province Natural Science Foundation","doi-asserted-by":"publisher","award":["262300421479"],"award-info":[{"award-number":["262300421479"]}],"id":[{"id":"10.13039\/501100006407","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004306","name":"Shandong Agricultural University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004306","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["U21A20260"],"award-info":[{"award-number":["U21A20260"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Veterinary Microbiology"],"published-print":{"date-parts":[[2026,5]]},"DOI":"10.1016\/j.vetmic.2026.110994","type":"journal-article","created":{"date-parts":[[2026,3,15]],"date-time":"2026-03-15T15:49:13Z","timestamp":1773589753000},"page":"110994","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":1,"special_numbering":"C","title":["Marek\u2019s disease virus UL16 tegument protein is essential for secondary envelopment and infectious virion formation"],"prefix":"10.1016","volume":"316","author":[{"given":"Xiuwen","family":"Yang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunzhe","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianhang","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingyu","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xing","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guoqing","family":"Zhuang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aijun","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.vetmic.2026.110994_bib1","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2022.109409","article-title":"Deletion of miR-M8 and miR-M13 eliminates the bursa atrophy induced by Marek\u2019s disease virus infection","volume":"268","author":"Bai","year":"2022","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib2","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.tim.2011.09.001","article-title":"Herpes simplex virus capsid assembly and DNA packaging: a present and future antiviral drug target","volume":"19","author":"Baines","year":"2011","journal-title":"Trends Microbiol"},{"key":"10.1016\/j.vetmic.2026.110994_bib3","doi-asserted-by":"crossref","first-page":"E1613","DOI":"10.1073\/pnas.1221896110","article-title":"Differential protein partitioning within the herpesvirus tegument and envelope underlies a complex and variable virion architecture","volume":"110","author":"Bohannon","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.vetmic.2026.110994_bib4","doi-asserted-by":"crossref","first-page":"10415","DOI":"10.1128\/JVI.05071-11","article-title":"Construction of a lytically replicating Kaposi's sarcoma-associated herpesvirus","volume":"85","author":"Budt","year":"2011","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib5","doi-asserted-by":"crossref","DOI":"10.1002\/cpps.34","article-title":"Preparation of cell cultures and vaccinia virus stocks","volume":"89","author":"Cotter","year":"2017","journal-title":"Curr. Protoc. Protein Sci."},{"key":"10.1016\/j.vetmic.2026.110994_bib6","doi-asserted-by":"crossref","DOI":"10.3389\/fmicb.2024.1431672","article-title":"The precise function of alphaherpesvirus tegument proteins and their interactions during the viral life cycle","volume":"15","author":"Cui","year":"2024","journal-title":"Front Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib7","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1038\/nature25438","article-title":"Structure and mutagenesis reveal essential capsid protein interactions for KSHV replication","volume":"553","author":"Dai","year":"2018","journal-title":"Nature"},{"key":"10.1016\/j.vetmic.2026.110994_bib8","doi-asserted-by":"crossref","first-page":"722","DOI":"10.1038\/nri3532","article-title":"Autophagy in infection, inflammation and immunity","volume":"13","author":"Deretic","year":"2013","journal-title":"Nat. Rev. Immunol."},{"key":"10.1016\/j.vetmic.2026.110994_bib9","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.00350-17","article-title":"The product of the herpes simplex virus 2 UL16 gene is critical for the egress of capsids from the nuclei of infected cells","volume":"91","author":"Gao","year":"2017","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib10","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.00629-18","article-title":"Comparative analysis of UL16 mutants derived from multiple strains of herpes simplex virus 2 (hsv-2) and hsv-1 reveals species-specific requirements for the UL16 protein","volume":"92","author":"Gao","year":"2018","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib11","doi-asserted-by":"crossref","DOI":"10.3390\/v13091828","article-title":"Disruption of the interaction between ORF33 and the conserved carboxyl-terminus of ORF45 abolishes progeny virion production of kaposi sarcoma-associated herpesvirus","volume":"13","author":"Gillen","year":"2021","journal-title":"Viruses"},{"key":"10.1016\/j.vetmic.2026.110994_bib12","doi-asserted-by":"crossref","first-page":"10582","DOI":"10.1128\/JVI.00497-09","article-title":"Open reading frame 33 of a gammaherpesvirus encodes a tegument protein essential for virion morphogenesis and egress","volume":"83","author":"Guo","year":"2009","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib13","doi-asserted-by":"crossref","first-page":"10582","DOI":"10.1128\/JVI.00497-09","article-title":"Open reading frame 33 of a gammaherpesvirus encodes a tegument protein essential for virion morphogenesis and egress","volume":"83","author":"Guo","year":"2009","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib14","doi-asserted-by":"crossref","first-page":"19798","DOI":"10.1073\/pnas.1212900109","article-title":"Function of glycoprotein E of herpes simplex virus requires coordinated assembly of three tegument proteins on its cytoplasmic tail","volume":"109","author":"Han","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.vetmic.2026.110994_bib15","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1007\/978-3-319-53168-7_6","article-title":"Herpesvirus capsid assembly and DNA packaging","volume":"223","author":"Heming","year":"2017","journal-title":"Adv. Anat. Embryol. Cell Biol."},{"key":"10.1016\/j.vetmic.2026.110994_bib16","doi-asserted-by":"crossref","DOI":"10.1016\/j.antiviral.2026.106344","article-title":"The UL23 thymidine kinase of Marek's disease virus is a target for anti-HSV drug development","volume":"247","author":"Kang","year":"2026","journal-title":"Antivir. Res"},{"key":"10.1016\/j.vetmic.2026.110994_bib17","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2026.110903","article-title":"UL24 deletion attenuates Marek's disease virus replication and pathogenicity","volume":"314","author":"Kang","year":"2026","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib18","article-title":"Calcium phosphate transfection","author":"Kingston","year":"2003","journal-title":"Curr. Protoc. Cell Biol. Chapter 20 Unit. 20. 23."},{"key":"10.1016\/j.vetmic.2026.110994_bib19","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":"10.1016\/j.vetmic.2026.110994_bib20","doi-asserted-by":"crossref","first-page":"1510","DOI":"10.1128\/JVI.79.3.1510-1522.2005","article-title":"Complex formation between the UL16 and UL21 tegument proteins of pseudorabies virus","volume":"79","author":"Klupp","year":"2005","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib21","doi-asserted-by":"crossref","DOI":"10.1128\/JVI.00295-18","article-title":"Varicella-zoster virus ORF9p binding to cellular adaptor protein complex 1 is important for viral infectivity","volume":"92","author":"Lebrun","year":"2018","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib22","doi-asserted-by":"crossref","first-page":"9254","DOI":"10.1128\/JVI.00443-11","article-title":"M94 is essential for the secondary envelopment of murine cytomegalovirus","volume":"85","author":"Maninger","year":"2011","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib23","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.virol.2009.12.004","article-title":"Complex mechanisms for the packaging of the UL16 tegument protein into herpes simplex virus","volume":"398","author":"Meckes","year":"2010","journal-title":"Virology"},{"key":"10.1016\/j.vetmic.2026.110994_bib24","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00810-20","article-title":"Conserved outer tegument component UL11 from herpes simplex virus 1 is an intrinsically disordered, rna-binding protein","volume":"11","author":"Metrick","year":"2020","journal-title":"mBio"},{"key":"10.1016\/j.vetmic.2026.110994_bib25","doi-asserted-by":"crossref","first-page":"7730","DOI":"10.1128\/JVI.02696-09","article-title":"A spread-deficient cytomegalovirus for assessment of first-target cells in vaccination","volume":"84","author":"Mohr","year":"2010","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib26","doi-asserted-by":"crossref","first-page":"5084","DOI":"10.3390\/v7092861","article-title":"Tegument assembly and secondary envelopment of alphaherpesviruses","volume":"7","author":"Owen","year":"2015","journal-title":"Viruses"},{"key":"10.1016\/j.vetmic.2026.110994_bib27","doi-asserted-by":"crossref","DOI":"10.3390\/ijms232113643","article-title":"Interplay between autophagy and herpes simplex Virus Type 1: ICP34.5, one of the main actors","volume":"23","author":"Ripa","year":"2022","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.vetmic.2026.110994_bib28","doi-asserted-by":"crossref","DOI":"10.1099\/jgv.0.002083","article-title":"The loss of both pUL16 and pUL21 in HSV-1-infected cells alters capsid-tegument composition, nuclear membrane architecture, cytoplasmic maturation and cell-to-cell spread","volume":"106","author":"Roddy","year":"2025","journal-title":"J. Gen. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib29","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1128\/JVI.02245-13","article-title":"Varicella-zoster virus ORF49 functions in the efficient production of progeny virus through its interaction with essential tegument protein ORF44","volume":"88","author":"Sadaoka","year":"2014","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib30","doi-asserted-by":"crossref","first-page":"5288","DOI":"10.1128\/JVI.00079-15","article-title":"Gammaherpesvirus tegument protein ORF33 is associated with intranuclear capsids at an early stage of the tegumentation process","volume":"89","author":"Shen","year":"2015","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib31","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1128\/JVI.02555-13","article-title":"Elucidation of the block to herpes simplex virus egress in the absence of tegument protein UL16 reveals a novel interaction with VP22","volume":"88","author":"Starkey","year":"2014","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib32","doi-asserted-by":"crossref","DOI":"10.3390\/v15102058","article-title":"Comprehensive analysis of the tegument proteins involved in capsid transport and virion morphogenesis of alpha, beta and gamma herpesviruses","volume":"15","author":"Sucharita","year":"2023","journal-title":"Viruses"},{"key":"10.1016\/j.vetmic.2026.110994_bib33","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1186\/s13567-021-00901-5","article-title":"UL28 and UL33 homologs of Marek's disease virus terminase complex involved in the regulation of cleavage and packaging of viral DNA are indispensable for replication in cultured cells","volume":"52","author":"Sun","year":"2021","journal-title":"Vet. Res"},{"key":"10.1016\/j.vetmic.2026.110994_bib34","doi-asserted-by":"crossref","DOI":"10.1128\/spectrum.02871-22","article-title":"Fully attenuated meq and pp38 double gene deletion mutant virus confers superior immunological protection against highly virulent marek's disease virus infection","volume":"10","author":"Sun","year":"2022","journal-title":"Microbiol. Spectr."},{"key":"10.1016\/j.vetmic.2026.110994_bib35","doi-asserted-by":"crossref","first-page":"191","DOI":"10.2144\/000112096","article-title":"Two-step red-mediated recombination for versatile high-efficiency markerless DNA manipulation in Escherichia coli","volume":"40","author":"Tischer","year":"2006","journal-title":"Biotechniques"},{"key":"10.1016\/j.vetmic.2026.110994_bib36","doi-asserted-by":"crossref","DOI":"10.1155\/2012\/472537","article-title":"Viral bacterial artificial chromosomes: generation, mutagenesis, and removal of mini-F sequences","volume":"2012","author":"Tischer","year":"2012","journal-title":"J. Biomed. Biotechnol."},{"key":"10.1016\/j.vetmic.2026.110994_bib37","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2024.110335","article-title":"MDV-encoded protein kinase U(S)3 phosphorylates WTAP to inhibit transcriptomic m(6)A modification and cellular protein translation","volume":"300","author":"Wang","year":"2025","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib38","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1128\/JVI.02738-15","article-title":"ORF33 and ORF38 of kaposi's sarcoma-associated herpesvirus interact and are required for optimal production of infectious progeny viruses","volume":"90","author":"Wu","year":"2016","journal-title":"J. Virol."},{"key":"10.1016\/j.vetmic.2026.110994_bib39","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2021.109327","article-title":"Pseudorabies virus UL16 protein influences the inhibition of LRPPRC for the viral proliferation","volume":"265","author":"Xu","year":"2022","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib40","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2021.109080","article-title":"Pseudorabies virus pUL16 assists the nuclear import of VP26 through protein-protein interaction","volume":"257","author":"Xu","year":"2021","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib41","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2021.109080","article-title":"Pseudorabies virus pUL16 assists the nuclear import of VP26 through protein-protein interaction","volume":"257","author":"Xu","year":"2021","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.vetmic.2026.110994_bib42","doi-asserted-by":"crossref","first-page":"9730","DOI":"10.1038\/srep09730","article-title":"Autophagy stimulation abrogates herpes simplex virus-1 infection","volume":"5","author":"Yakoub","year":"2015","journal-title":"Sci. Rep."},{"key":"10.1016\/j.vetmic.2026.110994_bib43","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijbiomac.2025.142730","article-title":"Unique long 11 and 16 protein act as a bridge for the unique long 11, 16, 21 protein and glycoprotein E interaction in duck plague virus replication","volume":"309","author":"Yang","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"10.1016\/j.vetmic.2026.110994_bib44","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2025.110622","article-title":"Marek's disease virus-encoded MicroRNA-M6-5p suppresses viral replication by targeting viral major capsid protein-coding gene UL19","volume":"307","author":"Zhou","year":"2025","journal-title":"Vet. Microbiol."}],"container-title":["Veterinary Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0378113526001252?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0378113526001252?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,4,27]],"date-time":"2026-04-27T14:32:46Z","timestamp":1777300366000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0378113526001252"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5]]},"references-count":44,"alternative-id":["S0378113526001252"],"URL":"https:\/\/doi.org\/10.1016\/j.vetmic.2026.110994","relation":{},"ISSN":["0378-1135"],"issn-type":[{"value":"0378-1135","type":"print"}],"subject":[],"published":{"date-parts":[[2026,5]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Marek\u2019s disease virus UL16 tegument protein is essential for secondary envelopment and infectious virion formation","name":"articletitle","label":"Article Title"},{"value":"Veterinary Microbiology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.vetmic.2026.110994","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 Elsevier B.V. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}],"article-number":"110994"}}