{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T02:36:07Z","timestamp":1760236567465,"version":"build-2065373602"},"reference-count":73,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2021,12,3]],"date-time":"2021-12-03T00:00:00Z","timestamp":1638489600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["1 R15 AI144683-01"],"award-info":[{"award-number":["1 R15 AI144683-01"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"name":"CMV Research Fund UTK","award":["RO1 1015 126"],"award-info":[{"award-number":["RO1 1015 126"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Pathogens"],"abstract":"<jats:p>Human cytomegalovirus (HCMV) is a ubiquitous pathogen infecting a majority of people worldwide, with diseases ranging from mild to life-threatening. Its clinical relevance in immunocompromised people and congenital infections have made treatment and vaccine development a top priority. Because of cytomegaloviruses\u2019 species specificity, murine cytomegalovirus (MCMV) models have historically informed and advanced translational CMV therapies. Using the phenomenon of centrifugal enhancement, we explored differences between MCMVs derived in vitro and in vivo. We found centrifugal enhancement on tissue culture-derived virus (TCV) was ~3\u00d7 greater compared with salivary gland derived virus (SGV). Using novel \u201cflow virometry\u201d, we found that TCV contained a distinct submicron particle composition compared to SGV. Using an inhibitor of exosome production, we show these submicron particles are not extracellular vesicles that contribute to centrifugal enhancement. We examined how these differences in submicron particles potentially contribute to differing centrifugal enhancement phenotypes, as well as broader in vivo vs. in vitro MCMV differences.<\/jats:p>","DOI":"10.3390\/pathogens10121577","type":"journal-article","created":{"date-parts":[[2021,12,6]],"date-time":"2021-12-06T03:10:38Z","timestamp":1638760238000},"page":"1577","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["MCMV Centrifugal Enhancement: A New Spin on an Old Topic"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3353-6042","authenticated-orcid":false,"given":"Trevor J.","family":"Hancock","sequence":"first","affiliation":[{"name":"Department of Microbiology, University of Tennessee, Knoxville, TN 37996, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4655-9427","authenticated-orcid":false,"given":"Morgan Lynn","family":"Hetzel","sequence":"additional","affiliation":[{"name":"Department of Microbiology, University of Tennessee, Knoxville, TN 37996, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andrea","family":"Ramirez","sequence":"additional","affiliation":[{"name":"Department of Microbiology, University of Tennessee, Knoxville, TN 37996, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tim E.","family":"Sparer","sequence":"additional","affiliation":[{"name":"Department of Microbiology, University of Tennessee, Knoxville, TN 37996, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2021,12,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1093\/cid\/ciu346","article-title":"Fields Virology, 6th Edition","volume":"59","author":"Liu","year":"2014","journal-title":"Clin. Infect. Dis."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/j.jcv.2007.11.014","article-title":"Human cytomegalovirus: Latency and reactivation in the myeloid lineage","volume":"41","author":"Sinclair","year":"2008","journal-title":"J. Clin. Virol."},{"key":"ref_3","first-page":"113","article-title":"Cytomegalovirus infection in patients with HIV infection","volume":"66","author":"Cheung","year":"1999","journal-title":"Mt. Sinai J. Med."},{"key":"ref_4","first-page":"36","article-title":"Congenital cytomegalovirus infection","volume":"1","author":"Demmler","year":"1994","journal-title":"Semin. Pediatr. Neurol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"260","DOI":"10.3947\/ic.2013.45.3.260","article-title":"Cytomegalovirus infections in solid organ transplantation: A review","volume":"45","author":"Ramanan","year":"2013","journal-title":"Infect. Chemother."},{"key":"ref_6","first-page":"65","article-title":"Congenital and perinatal cytomegalovirus infections: Clinical characteristics and pathogenic factors","volume":"20","author":"Stagno","year":"1984","journal-title":"Birth Defects Orig. Artic. Ser."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Reddehase, M.J., and Lemmermann, N.A.W. (2018). Mouse Model of Cytomegalovirus Disease and Immunotherapy in the Immunocompromised Host: Predictions for Medical Translation that Survived the \u201cTest of Time\u201d. Viruses, 10.","DOI":"10.3390\/v10120693"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1007\/s00430-015-0411-4","article-title":"Human cytomegalovirus: Taking the strain","volume":"204","author":"Wilkinson","year":"2015","journal-title":"Med. Microbiol. Immunol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"3191","DOI":"10.1172\/JCI42955","article-title":"Reconstruction of the complete human cytomegalovirus genome in a BAC reveals RL13 to be a potent inhibitor of replication","volume":"120","author":"Stanton","year":"2010","journal-title":"J. Clin. Investig."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"435","DOI":"10.3181\/00379727-86-21123","article-title":"Propagation of salivary gland virus of the mouse in tissue cultures","volume":"86","author":"Smith","year":"1954","journal-title":"Proc. Soc. Exp. Biol. Med."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/BF01317385","article-title":"Minor base sequence differences between the genomes of two strains of murine cytomegalovirus differing in virulence","volume":"64","author":"Misra","year":"1980","journal-title":"Arch. Virol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"6689","DOI":"10.1128\/JVI.00160-08","article-title":"Laboratory strains of murine cytomegalovirus are genetically similar to but phenotypically distinct from wild strains of virus","volume":"82","author":"Smith","year":"2008","journal-title":"J. Virol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"228","DOI":"10.1128\/iai.3.2.228-236.1971","article-title":"Virulence and attenuation of murine cytomegalovirus","volume":"3","author":"Osborn","year":"1971","journal-title":"Infect. Immun."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"840","DOI":"10.1128\/iai.33.3.840-847.1981","article-title":"Effects of cell source, mouse strain, and immunosuppressive treatment on production of virulent and attenuated murine cytomegalovirus","volume":"33","author":"Selgrade","year":"1981","journal-title":"Infect. Immun."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e00586-18","DOI":"10.1128\/mSphere.00586-18","article-title":"Anticytomegalovirus Peptides Point to New Insights for CMV Entry Mechanisms and the Limitations of In Vitro Screenings","volume":"4","author":"Jackson","year":"2019","journal-title":"mSphere"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1007\/BF01315153","article-title":"Neutralization of different strains of murine cytomegalovirus (MCMV)-effect of in vitro passage","volume":"69","author":"Chong","year":"1981","journal-title":"Arch. Virol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1016\/j.chom.2014.02.002","article-title":"Cytomegalovirus hijacks CX3CR1(hi) patrolling monocytes as immune-privileged vehicles for dissemination in mice","volume":"15","author":"Roback","year":"2014","journal-title":"Cell Host Microbe"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"5430","DOI":"10.1128\/jvi.64.11.5430-5440.1990","article-title":"Attenuated murine cytomegalovirus binds to N-acetylglucosamine, and shift to virulence may involve recognition of sialic acids","volume":"64","author":"Ravindranath","year":"1990","journal-title":"J. Virol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1016\/0042-6822(76)90325-1","article-title":"Properties of the multicapsid virions of murine cytomegalovirus","volume":"72","author":"Hudson","year":"1976","journal-title":"Virology"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1199","DOI":"10.1084\/jem.169.4.1199","article-title":"Site-restricted persistent cytomegalovirus infection after selective long-term depletion of CD4+ T lymphocytes","volume":"169","author":"Jonjic","year":"1989","journal-title":"J. Exp. Med."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1159\/000149701","article-title":"Studies on the morphogenesis of murine cytomegalovirus","volume":"26","author":"Weiland","year":"1986","journal-title":"Intervirology"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1128\/jvi.55.2.264-273.1985","article-title":"Interstitial murine cytomegalovirus pneumonia after irradiation: Characterization of cells that limit viral replication during established infection of the lungs","volume":"55","author":"Reddehase","year":"1985","journal-title":"J. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1128\/jvi.2.9.853-858.1968","article-title":"Enhancement of infectivity of murine cytomegalovirus in vitro by centrifugal inoculation","volume":"2","author":"Osborn","year":"1968","journal-title":"J. Virol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/0166-0934(88)90104-8","article-title":"Murine cytomegalovirus binds reversibly to mouse embryo fibroblasts: Implications for quantitation and explanation of centrifugal enhancement","volume":"22","author":"Hodgkin","year":"1988","journal-title":"J. Virol. Methods"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"431","DOI":"10.3181\/00379727-102-25275","article-title":"Plaque assay of Toxoplasma on monolayers of chick embryo fibroblasts","volume":"102","author":"Chaparas","year":"1959","journal-title":"Proc. Soc. Exp. Biol. Med."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1126\/science.131.3402.733","article-title":"Morphologic observations on trachoma virus in cell cultures","volume":"131","author":"Gordon","year":"1960","journal-title":"Science"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1128\/JVI.80.2.710-722.2006","article-title":"Human cytomegalovirus entry into epithelial and endothelial cells depends on genes UL128 to UL150 and occurs by endocytosis and low-pH fusion","volume":"80","author":"Ryckman","year":"2006","journal-title":"J. Virol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1016\/0042-6822(76)90326-3","article-title":"Cytomegalovirus infectivity: Analysis of the phenomenon of centrifugal enhancement of infectivity","volume":"72","author":"Hudson","year":"1976","journal-title":"Virology"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1016\/0166-0934(94)90047-7","article-title":"Enhancement of infectivity of hantavirus in cell culture by centrifugation","volume":"49","author":"Kariwa","year":"1994","journal-title":"J. Virol. Methods"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"10074","DOI":"10.1128\/JVI.74.21.10074-10080.2000","article-title":"Human immunodeficiency virus type 1 spinoculation enhances infection through virus binding","volume":"74","author":"Swiggard","year":"2000","journal-title":"J. Virol."},{"key":"ref_31","first-page":"2598","article-title":"Entry of human cytomegalovirus into retinal pigment epithelial and endothelial cells by endocytosis","volume":"40","author":"Bodaghi","year":"1999","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1128\/iai.21.1.281-285.1978","article-title":"Ultracentrifugal inoculation of herpes simplex virus","volume":"21","author":"Tenser","year":"1978","journal-title":"Infect. Immun."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1128\/iai.30.1.193-197.1980","article-title":"Mechanisms of herpes simplex virus infectivity enhanced by ultracentrifugal inoculation","volume":"30","author":"Tenser","year":"1980","journal-title":"Infect. Immun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/0166-0934(94)00119-2","article-title":"Enhancement of feline infectious peritonitis virus type I infection in cell cultures using low-speed centrifugation","volume":"51","author":"Hohdatsu","year":"1995","journal-title":"J. Virol. Methods"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Yan, R., Zhang, Y., Cai, D., Liu, Y., Cuconati, A., and Guo, H. (2015). Spinoculation Enhances HBV Infection in NTCP-Reconstituted Hepatocytes. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0129889"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/0042-6822(64)90265-X","article-title":"Human Cytomegalovirus. Morphology by Negative Staining","volume":"23","author":"Wright","year":"1964","journal-title":"Virology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e00609-20","DOI":"10.1128\/JVI.00609-20","article-title":"Human Cytomegalovirus Utilizes Extracellular Vesicles To Enhance Virus Spread","volume":"94","author":"Streck","year":"2020","journal-title":"J. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1038\/nrmicro.2017.60","article-title":"Viral effects on the content and function of extracellular vesicles","volume":"15","author":"Dittmer","year":"2017","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Urbanelli, L., Buratta, S., Tancini, B., Sagini, K., Delo, F., Porcellati, S., and Emiliani, C. (2019). The Role of Extracellular Vesicles in Viral Infection and Transmission. Vaccines, 7.","DOI":"10.3390\/vaccines7030102"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"12844","DOI":"10.1128\/JVI.05853-11","article-title":"Microvesicles and viral infection","volume":"85","author":"Meckes","year":"2011","journal-title":"J. Virol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"5200","DOI":"10.1128\/JVI.02470-14","article-title":"Exosomal communication goes viral","volume":"89","author":"Meckes","year":"2015","journal-title":"J. Virol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"e00088-18","DOI":"10.1128\/JVI.00088-18","article-title":"Role of Microvesicles in the Spread of Herpes Simplex Virus 1 in Oligodendrocytic Cells","volume":"92","author":"Praena","year":"2018","journal-title":"J. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2572","DOI":"10.3389\/fmicb.2018.02572","article-title":"Extracellular Vesicles in Herpes Viral Spread and Immune Evasion","volume":"9","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.virol.2018.08.008","article-title":"Human cytomegalovirus-infected cells release extracellular vesicles that carry viral surface proteins","volume":"524","author":"Zicari","year":"2018","journal-title":"Virology"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"9","DOI":"10.7554\/eLife.58288","article-title":"The host exosome pathway underpins biogenesis of the human cytomegalovirus virion","volume":"9","author":"Turner","year":"2020","journal-title":"eLife"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e01102-18","DOI":"10.1128\/JVI.01102-18","article-title":"Extracellular Vesicles Released by Herpes Simplex Virus 1-Infected Cells Block Virus Replication in Recipient Cells in a STING-Dependent Manner","volume":"92","author":"Deschamps","year":"2018","journal-title":"J. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"378","DOI":"10.1006\/viro.1997.8893","article-title":"The effect of herpes simplex virus type 1 L-particles on virus entry, replication, and the infectivity of naked herpesvirus DNA","volume":"239","author":"Dargan","year":"1997","journal-title":"Virology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1186\/s13578-020-00427-x","article-title":"Extracellular vesicles derived from different sources of mesenchymal stem cells: Therapeutic effects and translational potential","volume":"10","author":"Cai","year":"2020","journal-title":"Cell Biosci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1186\/s12964-020-00669-9","article-title":"Proteomics identifies differences in fibrotic potential of extracellular vesicles from human tendon and muscle fibroblasts","volume":"18","author":"Yeung","year":"2020","journal-title":"Cell Commun. Signal."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/0166-0934(88)90153-X","article-title":"Further studies on the mechanism of centrifugal enhancement of cytomegalovirus infectivity","volume":"19","author":"Hudson","year":"1988","journal-title":"J. Virol. Methods"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"4469","DOI":"10.1128\/JVI.02441-09","article-title":"The m74 gene product of murine cytomegalovirus (MCMV) is a functional homolog of human CMV gO and determines the entry pathway of MCMV","volume":"84","author":"Scrivano","year":"2010","journal-title":"J. Virol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1111\/tra.12355","article-title":"HCMV Induces Macropinocytosis for Host Cell Entry in Fibroblasts","volume":"17","author":"Hetzenecker","year":"2016","journal-title":"Traffic"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Wagner, F.M., Brizi\u0107, I., Prager, A., Tr\u0161an, T., Arapovi\u0107, M., Lemmermann, N.A.W., Podlech, J., Reddehase, M.J., Lemnitzer, F., and Bosse, J.B. (2013). The viral chemokine MCK-2 of murine cytomegalovirus promotes infection as part of a gH\/gL\/MCK-2 complex. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003493"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1182\/blood-2005-05-1833","article-title":"Cytomegalovirus MCK-2 controls mobilization and recruitment of myeloid progenitor cells to facilitate dissemination","volume":"107","author":"Noda","year":"2006","journal-title":"Blood"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"e01765-17","DOI":"10.1128\/JVI.01765-17","article-title":"Flow Virometry: A Powerful Tool To Functionally Characterize Viruses","volume":"92","author":"Lippe","year":"2018","journal-title":"J. Virol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"7432","DOI":"10.1038\/s41598-021-86688-z","article-title":"Flow virometry for process monitoring of live virus vaccines-lessons learned from ERVEBO","volume":"11","author":"Ricci","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1006\/viro.1993.1192","article-title":"Initiation of human cytomegalovirus infection requires initial interaction with cell surface heparan sulfate","volume":"193","author":"Compton","year":"1993","journal-title":"Virology"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Scrivano, L., Sinzger, C., Nitschko, H., Koszinowski, U.H., and Adler, B. (2011). HCMV spread and cell tropism are determined by distinct virus populations. PLoS Pathog., 7.","DOI":"10.1371\/journal.ppat.1001256"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"6243","DOI":"10.1128\/jvi.68.10.6243-6253.1994","article-title":"Peripheral blood mononuclear phagocytes mediate dissemination of murine cytomegalovirus","volume":"68","author":"Stoddart","year":"1994","journal-title":"J. Virol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1038\/s41419-020-02803-4","article-title":"Bioactive DNA from extracellular vesicles and particles","volume":"11","author":"Malkin","year":"2020","journal-title":"Cell Death Dis."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1038\/s41580-020-0251-y","article-title":"RNA delivery by extracellular vesicles in mammalian cells and its applications","volume":"21","author":"Breyne","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"2658","DOI":"10.1038\/s41467-018-05077-9","article-title":"Priming of dendritic cells by DNA-containing extracellular vesicles from activated T cells through antigen-driven contacts","volume":"9","author":"Torralba","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.4049\/jimmunol.1601694","article-title":"DNA-Containing Exosomes Derived from Cancer Cells Treated with Topotecan Activate a STING-Dependent Pathway and Reinforce Antitumor Immunity","volume":"198","author":"Kitai","year":"2017","journal-title":"J. Immunol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1470","DOI":"10.1038\/ncb1800","article-title":"Glioblastoma microvesicles transport RNA and proteins that promote tumour growth and provide diagnostic biomarkers","volume":"10","author":"Skog","year":"2008","journal-title":"Nat. Cell Biol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1038\/cr.2017.54","article-title":"Chemotherapy-induced intestinal inflammatory responses are mediated by exosome secretion of double-strand DNA via AIM2 inflammasome activation","volume":"27","author":"Lian","year":"2017","journal-title":"Cell Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.ebiom.2018.05.013","article-title":"Plasma Circulating Extracellular RNAs in Left Ventricular Remodeling Post-Myocardial Infarction","volume":"32","author":"Danielson","year":"2018","journal-title":"EBioMedicine"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1038\/nature12029","article-title":"A pathogenic picornavirus acquires an envelope by hijacking cellular membranes","volume":"496","author":"Feng","year":"2013","journal-title":"Nature"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"13109","DOI":"10.1073\/pnas.1221899110","article-title":"Exosome-mediated transmission of hepatitis C virus between human hepatoma Huh7.5 cells","volume":"110","author":"Ramakrishnaiah","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"682","DOI":"10.1016\/0042-6822(92)90906-6","article-title":"Noninfectious L-particles supply functions which can facilitate infection by HSV-1","volume":"190","author":"McLauchlan","year":"1992","journal-title":"Virology"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"11046","DOI":"10.1128\/JVI.01517-15","article-title":"L Particles Transmit Viral Proteins from Herpes Simplex Virus 1-Infected Mature Dendritic Cells to Uninfected Bystander Cells, Inducing CD83 Downmodulation","volume":"89","author":"Heilingloh","year":"2015","journal-title":"J. Virol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"17442","DOI":"10.1074\/jbc.M110.107821","article-title":"Secretory mechanisms and intercellular transfer of microRNAs in living cells","volume":"285","author":"Kosaka","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"2375","DOI":"10.1101\/gad.5.12b.2375","article-title":"p53 alteration is a common event in the spontaneous immortalization of primary BALB\/c murine embryo fibroblasts","volume":"5","author":"Harvey","year":"1991","journal-title":"Genes Dev."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"9966","DOI":"10.1128\/JVI.75.20.9966-9976.2001","article-title":"Murine cytomegalovirus CC chemokine homolog MCK-2 (m131-129) is a determinant of dissemination that increases inflammation at initial sites of infection","volume":"75","author":"Saederup","year":"2001","journal-title":"J. Virol."}],"container-title":["Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-0817\/10\/12\/1577\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:39:19Z","timestamp":1760168359000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-0817\/10\/12\/1577"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,12,3]]},"references-count":73,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["pathogens10121577"],"URL":"https:\/\/doi.org\/10.3390\/pathogens10121577","relation":{},"ISSN":["2076-0817"],"issn-type":[{"type":"electronic","value":"2076-0817"}],"subject":[],"published":{"date-parts":[[2021,12,3]]}}}