{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T00:08:14Z","timestamp":1790899694878,"version":"4.1.0"},"reference-count":73,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2015,3,4]],"date-time":"2015-03-04T00:00:00Z","timestamp":1425427200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"NIH","doi-asserted-by":"publisher","award":["R37 CA089441"],"award-info":[{"award-number":["R37 CA089441"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>Human endogenous retrovirus (HERV-K (HML-2)) proviruses are among the few endogenous retroviral elements in the human genome that retain coding sequence. HML-2 expression has been widely associated with human disease states, including different types of cancers as well as with HIV-1 infection. Understanding of the potential impact of this expression requires that it be annotated at the proviral level. Here, we utilized the high throughput capabilities of next-generation sequencing to profile HML-2 expression at the level of individual proviruses and secreted virions in the teratocarcinoma cell line Tera-1. We identified well-defined expression patterns, with transcripts emanating primarily from two proviruses located on chromosome 22, only one of which was efficiently packaged. Interestingly, there was a preference for transcripts of recently integrated proviruses, over those from other highly expressed but older elements, to be packaged into virions. We also assessed the promoter competence of the 5\u2019 long terminal repeats (LTRs) of expressed proviruses via a luciferase assay following transfection of Tera-1 cells. Consistent with the RNASeq results, we found that the activity of most LTRs corresponded to their transcript levels.<\/jats:p>","DOI":"10.3390\/v7030939","type":"journal-article","created":{"date-parts":[[2015,3,4]],"date-time":"2015-03-04T10:19:49Z","timestamp":1425464389000},"page":"939-968","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":73,"title":["Differential Expression of HERV-K (HML-2) Proviruses in Cells and Virions of the Teratocarcinoma Cell Line Tera-1"],"prefix":"10.3390","volume":"7","author":[{"given":"Neeru","family":"Bhardwaj","sequence":"first","affiliation":[{"name":"Graduate Program in Molecular Biology and Microbiology, Sackler School of Graduate Biomedical Sciences, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meagan","family":"Montesion","sequence":"additional","affiliation":[{"name":"Graduate Program in Genetics, Sackler School of Graduate Biomedical Sciences, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Farrah","family":"Roy","sequence":"additional","affiliation":[{"name":"Graduate Program in Genetics, Sackler School of Graduate Biomedical Sciences, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John","family":"Coffin","sequence":"additional","affiliation":[{"name":"Graduate Program in Molecular Biology and Microbiology, Sackler School of Graduate Biomedical Sciences, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"},{"name":"Graduate Program in Genetics, Sackler School of Graduate Biomedical Sciences, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"},{"name":"Department of Molecular Biology and Microbiology, Tufts University, 136 Harrison Avenue, Boston, MA 02111, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2015,3,4]]},"reference":[{"key":"ref_1","unstructured":"Coffin, J.M., Hughes, S.H., and Varmus, H.E. (1997). Retroviruses, Cold Spring Harbor Laboratory Press."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1146\/annurev.genom.7.080505.115700","article-title":"The Evolutionary Dynamics of Human Endogenous Retroviral Families","volume":"7","author":"Bannert","year":"2006","journal-title":"Annu. Rev. Genomics Hum. Genet."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1146\/annurev.genet.42.110807.091501","article-title":"Effects of Retroviruses on Host Genome Function","volume":"42","author":"Jern","year":"2008","journal-title":"Annu. Rev. Genet."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1668","DOI":"10.1073\/pnas.0307885100","article-title":"Human Endogenous Retrovirus K Solo-Ltr Formation and Insertional Polymorphisms: Implications for Human and Viral Evolution","volume":"101","author":"Hughes","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"e90","DOI":"10.1186\/1742-4690-8-90","article-title":"Identification, Characterization, and Comparative Genomic Distribution of the Herv-K (Hml-2) Group of Human Endogenous Retroviruses","volume":"8","author":"Subramanian","year":"2011","journal-title":"Retrovirology"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"12507","DOI":"10.1128\/JVI.79.19.12507-12514.2005","article-title":"Genomewide Screening Reveals High Levels of Insertional Polymorphism in the Human Endogenous Retrovirus Family Herv-K(Hml2): Implications for Present-Day Activity","volume":"79","author":"Belshaw","year":"2005","journal-title":"J. Virol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"9529","DOI":"10.1128\/JVI.00919-14","article-title":"Unfixed Endogenous Retroviral Insertions in the Human Population","volume":"88","author":"Marchi","year":"2014","journal-title":"J. Virol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1126\/science.2408338","article-title":"A New Class of Endogenous Human Retroviral Genomes","volume":"228","author":"Callahan","year":"1985","journal-title":"Science"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1203","DOI":"10.1128\/jvi.62.4.1203-1210.1988","article-title":"Expression of Human Sequences Related to Those of Mouse Mammary Tumor Virus","volume":"62","author":"Franklin","year":"1988","journal-title":"J. Virol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1126\/science.84.2172.162.a","article-title":"Some Possible Effects of Nursing on the Mammary Gland Tumor Incidence in Mice","volume":"84","author":"Bittner","year":"1936","journal-title":"Science"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1006\/viro.1993.1487","article-title":"Evidence That Herv-K Is the Endogenous Retrovirus Sequence That Codes for the Human Teratocarcinoma-Derived Retrovirus Htdv","volume":"196","author":"Boller","year":"1993","journal-title":"Virology"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1099\/0022-1317-82-3-591","article-title":"Phenotypic Heterogeneity of Human Endogenous Retrovirus Particles Produced by Teratocarcinoma Cell Lines","volume":"82","author":"Bieda","year":"2001","journal-title":"J. Gen. Virol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"10008","DOI":"10.1128\/JVI.01016-08","article-title":"Human Endogenous Retrovirus Family Herv-K(Hml-2) Rna Transcripts Are Selectively Packaged into Retroviral Particles Produced by the Human Germ Cell Tumor Line Tera-1 and Originate Mainly from a Provirus on Chromosome 22q11.21","volume":"82","author":"Ruprecht","year":"2008","journal-title":"J. Virol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1099\/0022-1317-65-5-887","article-title":"Human Teratocarcinomas Cultured in Vitro Produce Unique Retrovirus-Like Viruses","volume":"65","author":"Lower","year":"1984","journal-title":"J. Gen. Virol."},{"key":"ref_15","first-page":"293","article-title":"High Prevalence of Antibodies against Herv-K10 in Patients with Testicular Cancer but Not with Aids","volume":"8","author":"Goedert","year":"1999","journal-title":"Cancer Epidemiol. Biomark. Prev."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1128\/jvi.69.1.414-421.1995","article-title":"Human Endogenous Retrovirus K10: Expression of Gag Protein and Detection of Antibodies in Patients with Seminomas","volume":"69","author":"Sauter","year":"1995","journal-title":"J. Virol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1002\/ijc.11649","article-title":"Herv-K(Hml-2) Gag\/Env Antibodies as Indicator for Therapy Effect in Patients with Germ Cell Tumors","volume":"110","author":"Kleiman","year":"2004","journal-title":"Int. J. Cancer"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1002\/ijc.24902","article-title":"Beneficial and Detrimental Effects of Human Endogenous Retroviruses","volume":"126","author":"Kurth","year":"2010","journal-title":"Int. J. Cancer"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/S0014-5793(00)01197-2","article-title":"A Rev-Like Nes Mediates Cytoplasmic Localization of Herv-K Corf","volume":"468","author":"Boese","year":"2000","journal-title":"FEBS Lett."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"246","DOI":"10.3389\/fonc.2013.00246","article-title":"Herv-K(Hml-2), the Best Preserved Family of Hervs: Endogenization, Expression, and Implications in Health and Disease","volume":"3","author":"Hohn","year":"2013","journal-title":"Front. Oncol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2654","DOI":"10.1128\/jvi.70.4.2654-2657.1996","article-title":"Human Endogenous Retrovirus Expression and Reverse Transcriptase Activity in the T47d Mammary Carcinoma Cell Line","volume":"70","author":"Patience","year":"1996","journal-title":"J. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1006\/viro.1997.8670","article-title":"Type 1 Herv-K Genome Is Spliced into Subgenomic Transcripts in the Human Breast Tumor Cell Line T47d","volume":"234","author":"Etkind","year":"1997","journal-title":"Virology"},{"key":"ref_23","first-page":"1553","article-title":"Expression of Human Endogenous Retrovirus K Envelope Transcripts in Human Breast Cancer","volume":"7","author":"Frost","year":"2001","journal-title":"Clin. Cancer Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1038\/sj.onc.1206241","article-title":"Quantitation of Herv-K Env Gene Expression and Splicing in Human Breast Cancer","volume":"22","author":"Frost","year":"2003","journal-title":"Oncogene"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"5869","DOI":"10.1158\/0008-5472.CAN-07-6838","article-title":"Human Endogenous Retrovirus K Triggers an Antigen-Specific Immune Response in Breast Cancer Patients","volume":"68","author":"Radvanyi","year":"2008","journal-title":"Cancer Res."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1093\/jnci\/djr540","article-title":"Immunotherapeutic Potential of Anti-Human Endogenous Retrovirus-K Envelope Protein Antibodies in Targeting Breast Tumors","volume":"104","author":"Rycaj","year":"2012","journal-title":"J. Natl. Cancer Inst."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1097\/01.cmr.0000215031.07941.ca","article-title":"Expression of the Human Endogenous Retrovirus-K Transmembrane Envelope, Rec and Np9 Proteins in Melanomas and Melanoma Cell Lines","volume":"16","author":"Buscher","year":"2006","journal-title":"Melanoma Res."},{"key":"ref_28","first-page":"8735","article-title":"An Endogenous Retrovirus Derived from Human Melanoma Cells","volume":"63","author":"Muster","year":"2003","journal-title":"Cancer Res."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.jviromet.2006.03.029","article-title":"A New Real-Time-Rt-Pcr for Quantitation of Human Endogenous Retroviruses Type K (Herv-K) Rna Load in Plasma Samples: Increased Herv-K Rna Titers in Hiv-1 Patients with Haart Non-Suppressive Regimens","volume":"136","author":"Gonzalez","year":"2006","journal-title":"J. Virol. Methods"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1089\/aid.2006.22.979","article-title":"Detection of Herv-K(Hml-2) Viral Rna in Plasma of Hiv Type 1-Infected Individuals","volume":"22","author":"Kaplan","year":"2006","journal-title":"AIDS Res. Hum. Retrovir."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1089\/aid.2006.0117","article-title":"Hiv-1 Infection Increases the Expression of Human Endogenous Retroviruses Type K (Herv-K) in Vitro","volume":"23","author":"Lopez","year":"2007","journal-title":"AIDS Res. Hum. Retrovir."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e41021","DOI":"10.1371\/journal.pone.0041021","article-title":"Human Endogenous Retrovirus Expression is Inversely Associated with Chronic Immune Activation in Hiv-1 Infection","volume":"7","author":"Ormsby","year":"2012","journal-title":"PLoS One"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1128\/JVI.00602-11","article-title":"Characterization of Human Endogenous Retroviral Elements in the Blood of Hiv-1-Infected Individuals","volume":"86","author":"Kaplan","year":"2012","journal-title":"J. Virol."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Bhardwaj, N., Maldarelli, F., Mellors, J., and Coffin, J.M. (2014). HIV-1 Infection Leads to Increased Transcription of Herv-K (Hml-2) Proviruses in vivo but Not to Increased Virion Production. J. Virol., in press.","DOI":"10.1128\/JVI.01623-14"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1128\/JCM.43.1.406-413.2005","article-title":"Multiple, Linked Human Immunodeficiency Virus Type 1 Drug Resistance Mutations in Treatment-Experienced Patients Are Missed by Standard Genotype Analysis","volume":"43","author":"Palmer","year":"2005","journal-title":"J. Clin. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1186\/1742-4690-9-111","article-title":"Herv-H Rna is Abundant in Human Embryonic Stem Cells and a Precise Marker for Pluripotency","volume":"9","author":"Santoni","year":"2012","journal-title":"Retrovirology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"13837","DOI":"10.1128\/JVI.02388-13","article-title":"Comprehensive Analysis of Human Endogenous Retrovirus Group Herv-W Locus Transcription in Multiple Sclerosis Brain Lesions by High-Throughput Amplicon Sequencing","volume":"87","author":"Schmitt","year":"2013","journal-title":"J. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"6213","DOI":"10.1128\/JVI.00669-14","article-title":"Hiv-1 Interacts with Human Endogenous Retrovirus K (Hml-2) Envelopes Derived from Human Primary Lymphocytes","volume":"88","author":"Brinzevich","year":"2014","journal-title":"J. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Gonzalez-Hernandez, M.J., Cavalcoli, J.D., Sartor, M.A., Contreras-Galindo, R., Meng, F., Dai, M., Dube, D., Saha, A.K., Gitlin, S.D., and Omenn, G.S. (2014). Regulation of the Herv-K (Hml-2) Transcriptome by the HIV-1 Tat Protein. J. Virol.","DOI":"10.1128\/JVI.00556-14"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1006\/viro.1996.0443","article-title":"Endogenous D-Type (Herv-K) Related Sequences Are Packaged into Retroviral Particles in the Placenta and Possess Open Reading Frames for Reverse Transcriptase","volume":"222","author":"Simpson","year":"1996","journal-title":"Virology"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1006\/viro.1995.1205","article-title":"Restricted Expression of New Herv-K Members in Human Teratocarcinoma Cells","volume":"208","author":"Li","year":"1995","journal-title":"Virology"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: A Flexible Trimmer for Illumina Sequence Data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1038\/nmeth.1923","article-title":"Fast Gapped-Read Alignment with Bowtie 2","volume":"9","author":"Langmead","year":"2012","journal-title":"Nat. Methods"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"R36","DOI":"10.1186\/gb-2013-14-4-r36","article-title":"Tophat2: Accurate Alignment of Transcriptomes in the Presence of Insertions, Deletions and Gene Fusions","volume":"14","author":"Kim","year":"2013","journal-title":"Genome Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","article-title":"The Sequence Alignment\/Map Format and Samtools","volume":"25","author":"Li","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"R22","DOI":"10.1186\/gb-2011-12-3-r22","article-title":"Improving Rna-Seq Expression Estimates by Correcting for Fragment Bias","volume":"12","author":"Roberts","year":"2011","journal-title":"Genome Biol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1038\/nmeth.1226","article-title":"Mapping and Quantifying Mammalian Transcriptomes by Rna-Seq","volume":"5","author":"Mortazavi","year":"2008","journal-title":"Nat. Methods"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"D38","DOI":"10.1093\/nar\/gkq1172","article-title":"Database Resources of the National Center for Biotechnology Information","volume":"39","author":"Sayers","year":"2011","journal-title":"Nucleic Acids Res."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1093\/bib\/bbs017","article-title":"Integrative Genomics Viewer (Igv): High-Performance Genomics Data Visualization and Exploration","volume":"14","author":"Thorvaldsdottir","year":"2013","journal-title":"Brief. Bioinform."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"996","DOI":"10.1101\/gr.229102","article-title":"The Human Genome Browser at Ucsc","volume":"12","author":"Kent","year":"2002","journal-title":"Genome Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1093\/bioinformatics\/btr708","article-title":"Art: A Next-Generation Sequencing Read Simulator","volume":"28","author":"Huang","year":"2012","journal-title":"Bioinformatics"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"2725","DOI":"10.1093\/molbev\/mst197","article-title":"Mega6: Molecular Evolutionary Genetics Analysis Version 6.0","volume":"30","author":"Tamura","year":"2013","journal-title":"Mol. Biol. Evol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"D493","DOI":"10.1093\/nar\/gkh103","article-title":"The Ucsc Table Browser Data Retrieval Tool","volume":"32","author":"Karolchik","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"e115","DOI":"10.1093\/nar\/gks596","article-title":"Primer3\u2014New Capabilities and Interfaces","volume":"40","author":"Untergasser","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"8725","DOI":"10.1093\/nar\/15.21.8725","article-title":"A Novel Human Nonviral Retroposon Derived from an Endogenous Retrovirus","volume":"15","author":"Ono","year":"1987","journal-title":"Nucleic Acids Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"8762","DOI":"10.1128\/JVI.00751-08","article-title":"Hypermutation of an Ancient Human Retrovirus by Apobec3g","volume":"82","author":"Lee","year":"2008","journal-title":"J. Virol."},{"key":"ref_57","first-page":"1800","article-title":"A Novel Gene from the Human Endogenous Retrovirus K Expressed in Transformed Cells","volume":"8","author":"Armbruester","year":"2002","journal-title":"Clin. Cancer Res."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1128\/jvi.69.1.141-149.1995","article-title":"Identification of a Rev-Related Protein by Analysis of Spliced Transcripts of the Human Endogenous Retroviruses Htdv\/Herv-K","volume":"69","author":"Korbmacher","year":"1995","journal-title":"J. Virol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1915","DOI":"10.1101\/gad.17446611","article-title":"Integrative Annotation of Human Large Intergenic Noncoding Rnas Reveals Global Properties and Specific Subclasses","volume":"25","author":"Cabili","year":"2011","journal-title":"Genes Dev."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"19472","DOI":"10.1073\/pnas.1318172110","article-title":"Human-Specific Endogenous Retroviral Insert Serves as an Enhancer for the Schizophrenia-Linked Gene Prodh","volume":"110","author":"Suntsova","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.virusres.2004.02.036","article-title":"Tissue Specificity of Enhancer and Promoter Activities of a Herv-K(Hml-2) Ltr","volume":"104","author":"Ruda","year":"2004","journal-title":"Virus Res."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"e16","DOI":"10.1186\/1742-4690-10-16","article-title":"Endogenous Retrovirus-K Promoter: A Landing Strip for Inflammatory Transcription Factors?","volume":"10","author":"Manghera","year":"2013","journal-title":"Retrovirology"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"3436","DOI":"10.1128\/JVI.02539-10","article-title":"Expression of the Human Endogenous Retrovirus (Herv) Group Hml-2\/Herv-K Does Not Depend on Canonical Promoter Elements but Is Regulated by Transcription Factors Sp1 and Sp3","volume":"85","author":"Fuchs","year":"2011","journal-title":"J. Virol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1016\/j.virol.2005.11.007","article-title":"Functional Human Endogenous Retroviral Ltr Transcription Start Sites Are Located between the R and U5 Regions","volume":"346","author":"Kovalskaya","year":"2006","journal-title":"Virology"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"3366","DOI":"10.1007\/s00018-008-8496-1","article-title":"Endogenous Retroviruses and Cancer","volume":"65","author":"Ruprecht","year":"2008","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"e6","DOI":"10.1186\/1742-4690-9-6","article-title":"Hiv Infection and Herv Expression: A Review","volume":"9","year":"2012","journal-title":"Retrovirology"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1128\/JVI.79.1.341-352.2005","article-title":"Comprehensive Analysis of Human Endogenous Retrovirus Transcriptional Activity in Human Tissues with a Retrovirus-Specific Microarray","volume":"79","author":"Seifarth","year":"2005","journal-title":"J. Virol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"e354","DOI":"10.1186\/1471-2164-9-354","article-title":"Expression Patterns of Transcribed Human Endogenous Retrovirus Herv-K(Hml-2) Loci in Human Tissues and the Need for a Herv Transcriptome Project","volume":"9","author":"Flockerzi","year":"2008","journal-title":"BMC Genomics"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1128\/JVI.79.2.876-883.2005","article-title":"Cpg Methylation Directly Regulates Transcriptional Activity of the Human Endogenous Retrovirus Family Herv-K(Hml-2)","volume":"79","author":"Lavie","year":"2005","journal-title":"J. Virol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"e30","DOI":"10.1186\/1742-4690-8-30","article-title":"Identification of the Protease Cleavage Sites in a Reconstituted Gag Polyprotein of an Herv-K(Hml-2) Element","volume":"8","author":"George","year":"2011","journal-title":"Retrovirology"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"15573","DOI":"10.1128\/JVI.79.24.15573-15577.2005","article-title":"Identification of a Functional Envelope Protein from the Herv-K Family of Human Endogenous Retroviruses","volume":"79","author":"Dewannieux","year":"2005","journal-title":"J. Virol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1038\/nrmicro1210","article-title":"How Retroviruses Select Their Genomes","volume":"3","author":"Summers","year":"2005","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"6623","DOI":"10.1128\/JVI.02833-06","article-title":"Selective and Nonselective Packaging of Cellular Rnas in Retrovirus Particles","volume":"81","author":"Rulli","year":"2007","journal-title":"J. Virol."}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/7\/3\/939\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T20:43:08Z","timestamp":1760215388000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/7\/3\/939"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2015,3,4]]},"references-count":73,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2015,3]]}},"alternative-id":["v7030939"],"URL":"https:\/\/doi.org\/10.3390\/v7030939","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2015,3,4]]}}}