{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T23:04:57Z","timestamp":1787785497735,"version":"build-2784847793"},"reference-count":82,"publisher":"American Society of Hematology","issue":"5","license":[{"start":{"date-parts":[[2023,10,17]],"date-time":"2023-10-17T00:00:00Z","timestamp":1697500800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"content-domain":{"domain":["ashpublications.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2024,2,1]]},"abstract":"<jats:sec>\n                  <jats:title>Abstract<\/jats:title>\n                  <jats:p>Hematological malignancies such as Burkitt lymphoma (BL), Hodgkin lymphoma (HL), and diffuse large B-cell lymphoma (DLBCL) cause significant morbidity in humans. A substantial number of these lymphomas, particularly HL and DLBCLs have poorer prognosis because of their association with Epstein-Barr virus (EBV). Our earlier studies have shown that EBV-encoded nuclear antigen (EBNA2) upregulates programmed cell death ligand 1 in DLBCL and BLs by downregulating microRNA-34a. Here, we investigated whether EBNA2 affects the inducible costimulator (ICOS) ligand (ICOSL), a molecule required for efficient recognition of tumor cells by T cells through the engagement of ICOS on the latter. In virus-infected and EBNA2-transfected B-lymphoma cells, ICOSL expression was reduced. Our investigation of the molecular mechanisms revealed that this was due to an increase in microRNA-24 (miR-24) by EBNA2. By using ICOSL 3\u2032 untranslated region\u2013luciferase reporter system, we validated that ICOSL is an authentic miR-24 target. Transfection of anti\u2013miR-24 molecules in EBNA2-expressing lymphoma cells reconstituted ICOSL expression and increased tumor immunogenicity in mixed lymphocyte reactions. Because miR-24 is known to target c-MYC, an oncoprotein positively regulated by EBNA2, we analyzed its expression in anti\u2013miR-24 transfected lymphoma cells. Indeed, the reduction of miR-24 in EBNA2-expressing DLBCL further elevated c-MYC and increased apoptosis. Consistent with the in\u00a0vitro data, EBNA2-positive DLBCL biopsies expressed low ICOSL and high miR-24. We suggest that EBV evades host immune responses through EBNA2 by inducing miR-24 to reduce ICOSL expression, and for simultaneous rheostatic maintenance of proproliferative c-MYC levels. Overall, these data identify miR-24 as a potential therapeutically relevant target in EBV-associated lymphomas.<\/jats:p>\n               <\/jats:sec>","DOI":"10.1182\/blood.2023021346","type":"journal-article","created":{"date-parts":[[2023,10,17]],"date-time":"2023-10-17T19:01:07Z","timestamp":1697569267000},"page":"429-443","update-policy":"https:\/\/doi.org\/10.1182\/blood.2019cm0000","source":"Crossref","is-referenced-by-count":27,"title":["Epstein-Barr virus\u2013encoded EBNA2 downregulates ICOSL by inducing miR-24 in B-cell lymphoma"],"prefix":"10.1182","volume":"143","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9258-4191","authenticated-orcid":false,"given":"Martina","family":"Leopizzi","sequence":"first","affiliation":[{"name":"1Department of Medico-surgical Sciences and Biotechnologies, Sapienza University, Latina, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lucia","family":"Mundo","sequence":"additional","affiliation":[{"name":"2Department of Medical Biotechnology, University of Siena, Siena, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7182-2883","authenticated-orcid":false,"given":"Elena","family":"Messina","sequence":"additional","affiliation":[{"name":"3Department of Experimental Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9290-874X","authenticated-orcid":false,"given":"Federica","family":"Campolo","sequence":"additional","affiliation":[{"name":"3Department of Experimental Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefano","family":"Lazzi","sequence":"additional","affiliation":[{"name":"2Department of Medical Biotechnology, University of Siena, Siena, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Antonio","family":"Angeloni","sequence":"additional","affiliation":[{"name":"3Department of Experimental Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9280-8917","authenticated-orcid":false,"given":"Cinzia","family":"Marchese","sequence":"additional","affiliation":[{"name":"3Department of Experimental Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lorenzo","family":"Leoncini","sequence":"additional","affiliation":[{"name":"2Department of Medical Biotechnology, University of Siena, Siena, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3716-4303","authenticated-orcid":false,"given":"Carla","family":"Giordano","sequence":"additional","affiliation":[{"name":"4Department of Radiology, Oncology and Pathology, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Frank","family":"Slack","sequence":"additional","affiliation":[{"name":"5Department of Pathology, Beth Israel Deaconess Medical Center, Harvard Medical School, Boston, MA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9574-6304","authenticated-orcid":false,"given":"Pankaj","family":"Trivedi","sequence":"additional","affiliation":[{"name":"3Department of Experimental Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0212-6734","authenticated-orcid":false,"given":"Eleni","family":"Anastasiadou","sequence":"additional","affiliation":[{"name":"6Department of Clinical and Molecular Medicine, Sapienza University, Rome, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"234","reference":[{"issue":"1","key":"2024020116214416200_bib1","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1007\/s00432-021-03824-y","article-title":"Estimating the global burden of Epstein-Barr virus-related cancers","volume":"148","author":"Wong","year":"2022","journal-title":"J\u00a0Cancer Res Clin Oncol"},{"key":"2024020116214416200_bib2","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1146\/annurev.med.56.082103.104727","article-title":"Post-transplant lymphoproliferative disorders","volume":"56","author":"Gottschalk","year":"2005","journal-title":"Annu Rev Med"},{"issue":"7","key":"2024020116214416200_bib3","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1182\/blood.2020005469","article-title":"Hematologic cancers in individuals infected by HIV","volume":"139","author":"Carbone","year":"2022","journal-title":"Blood"},{"issue":"12","key":"2024020116214416200_bib4","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1038\/nrc.2016.92","article-title":"Epstein-Barr virus: more than 50 years old and still providing surprises","volume":"16","author":"Young","year":"2016","journal-title":"Nat Rev Cancer"},{"issue":"Pt 1","key":"2024020116214416200_bib5","first-page":"17","article-title":"Tumor associations of EBV--historical perspectives","volume":"390","author":"Klein","year":"2015","journal-title":"Curr Top Microbiol Immunol"},{"issue":"4","key":"2024020116214416200_bib6","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1002\/ajh.25760","article-title":"EBV-positive diffuse large B-cell lymphoma, not otherwise specified: 2020 update on diagnosis, risk-stratification and management","volume":"95","author":"Beltran","year":"2020","journal-title":"Am J Hematol"},{"issue":"6769","key":"2024020116214416200_bib7","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1038\/35000501","article-title":"Distinct types of diffuse large B-cell lymphoma identified by gene expression profiling","volume":"403","author":"Alizadeh","year":"2000","journal-title":"Nature"},{"key":"2024020116214416200_bib8","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1177\/117693510700300004","article-title":"Germinal center B cell-like (GCB) and activated B cell-like (ABC) type of diffuse large B cell lymphoma (DLBCL): analysis of molecular predictors, signatures, cell cycle state and patient survival","volume":"3","author":"Blenk","year":"2007","journal-title":"Cancer Inform"},{"issue":"4","key":"2024020116214416200_bib9","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/j.ccell.2020.03.015","article-title":"A\u00a0probabilistic classification tool for genetic subtypes of diffuse large B cell lymphoma with therapeutic implications","volume":"37","author":"Wright","year":"2020","journal-title":"Cancer Cell"},{"issue":"12","key":"2024020116214416200_bib10","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1111\/j.1349-7006.2000.tb00909.x","article-title":"The characteristics of Epstein-Barr virus (EBV)-positive diffuse large B-cell lymphoma: comparison between EBV(+) and EBV(-) cases in Japanese population","volume":"91","author":"Kuze","year":"2000","journal-title":"Jpn J Cancer Res"},{"issue":"1","key":"2024020116214416200_bib11","doi-asserted-by":"crossref","first-page":"39","DOI":"10.3109\/10428194.2015.1040014","article-title":"In Europe expression of EBNA2 is associated with poor survival in EBV-positive diffuse large B-cell lymphoma of the elderly","volume":"57","author":"Stuhlmann-Laeisz","year":"2016","journal-title":"Leuk Lymphoma"},{"issue":"11","key":"2024020116214416200_bib12","doi-asserted-by":"crossref","first-page":"1468","DOI":"10.1182\/blood.2020008520","article-title":"EBV-associated primary CNS lymphoma occurring after immunosuppression is a distinct immunobiological entity","volume":"137","author":"Gandhi","year":"2021","journal-title":"Blood"},{"issue":"3","key":"2024020116214416200_bib13","doi-asserted-by":"crossref","first-page":"5361","DOI":"10.18632\/oncotarget.14243","article-title":"Epstein-Barr virus EBNA2 directs doxorubicin resistance of B cell lymphoma through CCL3 and CCL4-mediated activation of NF-\u03baB and Btk","volume":"8","author":"Kim","year":"2017","journal-title":"Oncotarget"},{"issue":"1","key":"2024020116214416200_bib14","doi-asserted-by":"crossref","first-page":"e131","DOI":"10.1038\/emm.2014.84","article-title":"Epstein-Barr virus latent genes","volume":"47","author":"Kang","year":"2015","journal-title":"Exp Mol Med"},{"issue":"10","key":"2024020116214416200_bib15","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1038\/nri1201","article-title":"B\u00a0cells under influence: transformation of B cells by Epstein-Barr virus","volume":"3","author":"Kuppers","year":"2003","journal-title":"Nat Rev Immunol"},{"issue":"6","key":"2024020116214416200_bib16","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/j.semcancer.2009.07.004","article-title":"Burkitt's lymphoma: the Rosetta Stone deciphering Epstein-Barr virus biology","volume":"19","author":"Rowe","year":"2009","journal-title":"Semin Cancer Biol"},{"issue":"23","key":"2024020116214416200_bib17","doi-asserted-by":"crossref","first-page":"9558","DOI":"10.1073\/pnas.86.23.9558","article-title":"Epstein-Barr virus nuclear protein 2 is a key determinant of lymphocyte transformation","volume":"86","author":"Cohen","year":"1989","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"2024020116214416200_bib18","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1128\/jvi.44.3.834-844.1982","article-title":"Non-immortalizing P3J-HR-1 Epstein-Barr virus: a deletion mutant of its transforming parent, Jijoye","volume":"44","author":"Rabson","year":"1982","journal-title":"J\u00a0Virol"},{"issue":"16","key":"2024020116214416200_bib19","doi-asserted-by":"crossref","first-page":"7568","DOI":"10.1073\/pnas.91.16.7568","article-title":"The Epstein-Barr virus nuclear antigen 2 transactivator is directed to response elements by the J kappa recombination signal binding protein","volume":"91","author":"Grossman","year":"1994","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"2024020116214416200_bib20","doi-asserted-by":"crossref","first-page":"e1005339","DOI":"10.1371\/journal.ppat.1005339","article-title":"EBNA2 drives formation of new chromosome binding sites and target genes for B-cell master regulatory transcription factors RBP-jkappa and EBF1","volume":"12","author":"Lu","year":"2016","journal-title":"PLoS Pathog"},{"issue":"5","key":"2024020116214416200_bib21","doi-asserted-by":"crossref","first-page":"4481","DOI":"10.1128\/JVI.73.5.4481-4484.1999","article-title":"The proto-oncogene c-myc is a direct target gene of Epstein-Barr virus nuclear antigen 2","volume":"73","author":"Kaiser","year":"1999","journal-title":"J\u00a0Virol"},{"issue":"1","key":"2024020116214416200_bib22","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1038\/nrc.2017.99","article-title":"Non-coding RNA networks in cancer","volume":"18","author":"Anastasiadou","year":"2018","journal-title":"Nat Rev Cancer"},{"key":"2024020116214416200_bib23","doi-asserted-by":"crossref","first-page":"101272","DOI":"10.1016\/j.coviro.2022.101272","article-title":"MicroRNA-mediated control of Epstein-Barr virus infection and potential diagnostic and therapeutic implications","volume":"56","author":"Skalsky","year":"2022","journal-title":"Curr Opin Virol"},{"issue":"42","key":"2024020116214416200_bib24","doi-asserted-by":"crossref","first-page":"E6467","DOI":"10.1073\/pnas.1605884113","article-title":"Epstein-Barr virus microRNAs reduce immune surveillance by virus-specific CD8+ T cells","volume":"113","author":"Albanese","year":"2016","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"2024020116214416200_bib25","doi-asserted-by":"crossref","first-page":"866","DOI":"10.1038\/s41467-022-28479-2","article-title":"EBV miRNAs BART11 and BART17-3p promote immune escape through the enhancer-mediated transcription of PD-L1","volume":"13","author":"Wang","year":"2022","journal-title":"Nat Commun"},{"issue":"25","key":"2024020116214416200_bib26","doi-asserted-by":"crossref","first-page":"2261","DOI":"10.1182\/blood.2019000889","article-title":"EBV microRNA-BHRF1-2-5p targets the 3'UTR of immune checkpoint ligands PD-L1 and PD-L2","volume":"134","author":"Cristino","year":"2019","journal-title":"Blood"},{"issue":"11","key":"2024020116214416200_bib27","doi-asserted-by":"crossref","first-page":"2343","DOI":"10.1038\/leu.2012.108","article-title":"Differential regulation of miR-21 and miR-146a by Epstein-Barr virus-encoded EBNA2","volume":"26","author":"Rosato","year":"2012","journal-title":"Leukemia"},{"issue":"22","key":"2024020116214416200_bib28","doi-asserted-by":"crossref","first-page":"11670","DOI":"10.1128\/JVI.01248-10","article-title":"Virally induced cellular microRNA miR-155 plays a key role in B-cell immortalization by Epstein-Barr virus","volume":"84","author":"Linnstaedt","year":"2010","journal-title":"J\u00a0Virol"},{"issue":"14-15","key":"2024020116214416200_bib29","doi-asserted-by":"crossref","first-page":"1156","DOI":"10.1016\/j.micinf.2011.07.007","article-title":"The role of microRNAs in Epstein-Barr virus latency and lytic reactivation","volume":"13","author":"Forte","year":"2011","journal-title":"Microbes Infect"},{"issue":"22","key":"2024020116214416200_bib30","doi-asserted-by":"crossref","first-page":"3595","DOI":"10.4161\/cc.7.22.7120","article-title":"Epstein-Barr virus-mediated dysregulation of human microRNA expression","volume":"7","author":"Godshalk","year":"2008","journal-title":"Cell Cycle"},{"issue":"2","key":"2024020116214416200_bib31","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.cell.2015.03.030","article-title":"Immune checkpoint targeting in cancer therapy: toward combination strategies with curative potential","volume":"161","author":"Sharma","year":"2015","journal-title":"Cell"},{"key":"2024020116214416200_bib32","doi-asserted-by":"crossref","first-page":"101480","DOI":"10.1016\/j.smim.2021.101480","article-title":"Emerging concepts in PD-1 checkpoint biology","volume":"52","author":"Pauken","year":"2021","journal-title":"Semin Immunol"},{"issue":"8","key":"2024020116214416200_bib33","doi-asserted-by":"crossref","first-page":"1652","DOI":"10.1016\/j.cell.2023.03.006","article-title":"Immune checkpoint therapy-current perspectives and future directions","volume":"186","author":"Sharma","year":"2023","journal-title":"Cell"},{"issue":"92","key":"2024020116214416200_bib34","doi-asserted-by":"crossref","first-page":"36411","DOI":"10.18632\/oncotarget.26381","article-title":"Epstein-Barr virus: from kisses to cancer, an ingenious immune evader","volume":"9","author":"Trivedi","year":"2018","journal-title":"Oncotarget"},{"issue":"13","key":"2024020116214416200_bib35","doi-asserted-by":"crossref","first-page":"4567","DOI":"10.1073\/pnas.84.13.4567","article-title":"Down-regulation of class I HLA antigens and of the Epstein-Barr virus-encoded latent membrane protein in Burkitt lymphoma lines","volume":"84","author":"Masucci","year":"1987","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5104","key":"2024020116214416200_bib36","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1126\/science.7682013","article-title":"HLA-A11 epitope loss isolates of Epstein-Barr virus from a highly A11+ population","volume":"260","author":"de Campos-Lima","year":"1993","journal-title":"Science"},{"issue":"5361","key":"2024020116214416200_bib37","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1126\/science.280.5361.248","article-title":"Viral strategies of immune evasion","volume":"280","author":"Ploegh","year":"1998","journal-title":"Science"},{"issue":"1","key":"2024020116214416200_bib38","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1038\/s41375-018-0178-x","article-title":"Epstein-Barr virus-encoded EBNA2 alters immune checkpoint PD-L1 expression by downregulating miR-34a in B-cell lymphomas","volume":"33","author":"Anastasiadou","year":"2019","journal-title":"Leukemia"},{"issue":"11","key":"2024020116214416200_bib39","doi-asserted-by":"crossref","first-page":"2179","DOI":"10.1080\/1042819021000032917","article-title":"A\u00a0novel B-cell line (U-2932) established from a patient with diffuse large B-cell lymphoma following Hodgkin lymphoma","volume":"43","author":"Amini","year":"2002","journal-title":"Leuk Lymphoma"},{"issue":"5","key":"2024020116214416200_bib40","doi-asserted-by":"crossref","first-page":"2274","DOI":"10.1128\/JVI.01822-06","article-title":"EBNA2 interferes with the germinal center phenotype by downregulating BCL6 and TCL1 in non-Hodgkin's lymphoma cells","volume":"81","author":"Boccellato","year":"2007","journal-title":"J\u00a0Virol"},{"issue":"2","key":"2024020116214416200_bib41","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1002\/ijc.2910390215","article-title":"Expression of B-cell-specific markers in different Burkitt lymphoma subgroups","volume":"39","author":"Ehlin-Henriksson","year":"1987","journal-title":"Int J Cancer"},{"issue":"4","key":"2024020116214416200_bib42","first-page":"276","article-title":"Establishment and characterization of an Epstein-Barr virus (EBC)-negative lymphoblastoid B cell line (BJA-B) from an exceptional, EBV-genome-negative African Burkitt's lymphoma","volume":"22","author":"Menezes","year":"1975","journal-title":"Biomedicine"},{"issue":"2","key":"2024020116214416200_bib43","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1002\/ijc.2910400213","article-title":"Conversion of the lymphoma line \"BJAB\" by Epstein-Barr virus into phenotypically altered sublines is accompanied by increased c-myc mRNA levels","volume":"40","author":"Wennborg","year":"1987","journal-title":"Int J Cancer"},{"issue":"2","key":"2024020116214416200_bib44","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1006\/viro.1995.0084","article-title":"Epstein-Barr virus nuclear antigen 2-estrogen receptor fusion proteins transactivate viral and cellular genes and interact with RBP-J kappa in a conditional fashion","volume":"214","author":"Kempkes","year":"1995","journal-title":"Virology"},{"issue":"3","key":"2024020116214416200_bib45","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1128\/jvi.43.3.952-968.1982","article-title":"Deletion of the nontransforming Epstein-Barr virus strain P3HR-1 causes fusion of the large internal repeat to the DSL region","volume":"43","author":"Bornkamm","year":"1982","journal-title":"J\u00a0Virol"},{"issue":"5","key":"2024020116214416200_bib46","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1002\/ijc.2910400521","article-title":"Paired Epstein-Barr virus (EBV)-negative and EBV-converted Burkitt lymphoma lines: stimulatory capacity in allogeneic mixed lymphocyte cultures","volume":"40","author":"Avila-Cari\u00f1o","year":"1987","journal-title":"Int J Cancer"},{"key":"2024020116214416200_bib47","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1186\/1471-2105-11-249","article-title":"Gene bi-targeting by viral and human miRNAs","volume":"11","author":"Veksler-Lublinsky","year":"2010","journal-title":"BMC Bioinformatics"},{"key":"2024020116214416200_bib48","doi-asserted-by":"crossref","first-page":"e05005","DOI":"10.7554\/eLife.05005","article-title":"Predicting effective microRNA target sites in mammalian mRNAs","volume":"4","author":"Agarwal","year":"2015","journal-title":"Elife"},{"issue":"10","key":"2024020116214416200_bib49","doi-asserted-by":"crossref","first-page":"e0206239","DOI":"10.1371\/journal.pone.0206239","article-title":"miRWalk: an online resource for prediction of microRNA binding sites","volume":"13","author":"Sticht","year":"2018","journal-title":"PLoS One"},{"issue":"1","key":"2024020116214416200_bib50","doi-asserted-by":"crossref","first-page":"24","DOI":"10.3390\/microarrays2010024","article-title":"Profiling pre-microRNA and mature microRNA expressions using a single microarray and avoiding separate sample preparation","volume":"2","author":"Gan","year":"2013","journal-title":"Microarrays (Basel)"},{"issue":"1","key":"2024020116214416200_bib51","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.ymeth.2007.09.006","article-title":"Real-time PCR quantification of precursor and mature microRNA","volume":"44","author":"Schmittgen","year":"2008","journal-title":"Methods"},{"issue":"5","key":"2024020116214416200_bib52","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1016\/j.molcel.2009.08.020","article-title":"miR-24 Inhibits cell proliferation by targeting E2F2, MYC, and other cell-cycle genes via binding to \"seedless\" 3'UTR microRNA recognition elements","volume":"35","author":"Lal","year":"2009","journal-title":"Mol Cell"},{"issue":"1","key":"2024020116214416200_bib53","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/0092-8674(92)90123-T","article-title":"Induction of apoptosis in fibroblasts by c-myc protein","volume":"69","author":"Evan","year":"1992","journal-title":"Cell"},{"issue":"7","key":"2024020116214416200_bib54","doi-asserted-by":"crossref","first-page":"1720","DOI":"10.1038\/s41375-022-01620-2","article-title":"The 5th edition of the World Health Organization classification of haematolymphoid tumours: lymphoid neoplasms","volume":"36","author":"Alaggio","year":"2022","journal-title":"Leukemia"},{"issue":"1","key":"2024020116214416200_bib55","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1182\/blood-2003-05-1545","article-title":"Confirmation of the molecular classification of diffuse large B-cell lymphoma by immunohistochemistry using a tissue microarray","volume":"103","author":"Hans","year":"2004","journal-title":"Blood"},{"issue":"1","key":"2024020116214416200_bib82","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1186\/s13046-023-02752-8","article-title":"The new world of RNA diagnostics and therapeutics","volume":"42","author":"Blandino","year":"2023","journal-title":"J Exp Clin Cancer Res"},{"issue":"1","key":"2024020116214416200_bib56","doi-asserted-by":"crossref","first-page":"e000544","DOI":"10.1136\/esmoopen-2019-000544","article-title":"The rationale behind targeting the ICOS-ICOS ligand costimulatory pathway in cancer immunotherapy","volume":"5","author":"Solinas","year":"2020","journal-title":"ESMO Open"},{"issue":"16","key":"2024020116214416200_bib57","doi-asserted-by":"crossref","first-page":"5445","DOI":"10.1158\/0008-5472.CAN-11-1138","article-title":"The ICOS\/ICOSL pathway is required for optimal antitumor responses mediated by anti-CTLA-4 therapy","volume":"71","author":"Fu","year":"2011","journal-title":"Cancer Res"},{"issue":"4","key":"2024020116214416200_bib58","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1084\/jem.20130590","article-title":"Engagement of the ICOS pathway markedly enhances efficacy of CTLA-4 blockade in cancer immunotherapy","volume":"211","author":"Fan","year":"2014","journal-title":"J\u00a0Exp Med"},{"issue":"1","key":"2024020116214416200_bib59","doi-asserted-by":"crossref","first-page":"38","DOI":"10.3390\/biom12010038","article-title":"Co-stimulatory molecules during immune control of Epstein Barr virus infection","volume":"12","author":"M\u00fcnz","year":"2021","journal-title":"Biomolecules"},{"issue":"12","key":"2024020116214416200_bib60","doi-asserted-by":"crossref","first-page":"2185","DOI":"10.1101\/gr.264705.120","article-title":"Epstein-Barr virus nuclear antigen 2 extensively rewires the human chromatin landscape at autoimmune risk loci","volume":"31","author":"Hong","year":"2021","journal-title":"Genome Res"},{"issue":"6","key":"2024020116214416200_bib61","doi-asserted-by":"crossref","first-page":"1491","DOI":"10.1002\/ijc.29489","article-title":"Epstein-Barr virus infection induces miR-21 in terminally differentiated malignant B cells","volume":"137","author":"Anastasiadou","year":"2015","journal-title":"Int J Cancer"},{"issue":"2","key":"2024020116214416200_bib62","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1016\/j.virol.2008.09.018","article-title":"Epstein-Barr virus growth\/latency III program alters cellular microRNA expression","volume":"382","author":"Cameron","year":"2008","journal-title":"Virology"},{"issue":"5","key":"2024020116214416200_bib63","first-page":"5769","article-title":"Expression and significance of miR-24 and miR-101 in patients with advanced gastric cancer","volume":"16","author":"Dong","year":"2018","journal-title":"Oncol Lett"},{"issue":"3","key":"2024020116214416200_bib64","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1016\/j.oraloncology.2009.12.005","article-title":"miR-24 up-regulation in oral carcinoma: positive association from clinical and in\u00a0vitro analysis","volume":"46","author":"Lin","year":"2010","journal-title":"Oral Oncol"},{"issue":"1","key":"2024020116214416200_bib65","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1186\/s12864-022-09018-1","article-title":"Comprehensive microRNA analysis across genome-edited colorectal cancer organoid models reveals miR-24 as a candidate regulator of cell survival","volume":"23","author":"Villanueva","year":"2022","journal-title":"BMC Genomics"},{"issue":"7","key":"2024020116214416200_bib66","doi-asserted-by":"crossref","first-page":"e2557","DOI":"10.1371\/journal.pone.0002557","article-title":"Aberrant expression of oncogenic and tumor-suppressive microRNAs in cervical cancer is required for cancer cell growth","volume":"3","author":"Wang","year":"2008","journal-title":"PLoS One"},{"issue":"8","key":"2024020116214416200_bib67","doi-asserted-by":"crossref","first-page":"2128","DOI":"10.1099\/jgv.0.000870","article-title":"Antibodies against the mono-methylated arginine-glycine repeat (MMA-RG) of the Epstein-Barr virus nuclear antigen 2 (EBNA2) identify potential cellular proteins targeted in viral transformation","volume":"98","author":"Ayoubian","year":"2017","journal-title":"J\u00a0Gen Virol"},{"issue":"7014","key":"2024020116214416200_bib68","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1038\/nature03120","article-title":"The Microprocessor complex mediates the genesis of microRNAs","volume":"432","author":"Gregory","year":"2004","journal-title":"Nature"},{"issue":"20","key":"2024020116214416200_bib69","doi-asserted-by":"crossref","first-page":"4196","DOI":"10.1016\/j.yexcr.2007.07.020","article-title":"Nucleolar localization of DGCR8 and identification of eleven DGCR8-associated proteins","volume":"313","author":"Shiohama","year":"2007","journal-title":"Exp Cell Res"},{"issue":"30","key":"2024020116214416200_bib70","doi-asserted-by":"crossref","first-page":"e2200512119","DOI":"10.1073\/pnas.2200512119","article-title":"EBNA2-EBF1 complexes promote MYC expression and metabolic processes driving S-phase progression of Epstein-Barr virus-infected B cells","volume":"119","author":"Beer","year":"2022","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"6","key":"2024020116214416200_bib71","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.ccr.2008.10.018","article-title":"Distinct thresholds govern Myc's biological output in\u00a0vivo","volume":"14","author":"Murphy","year":"2008","journal-title":"Cancer Cell"},{"issue":"3","key":"2024020116214416200_bib72","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1016\/S0092-8674(02)00738-9","article-title":"Suppression of Myc-induced apoptosis in beta cells exposes multiple oncogenic properties of Myc and triggers carcinogenic progression","volume":"109","author":"Pelengaris","year":"2002","journal-title":"Cell"},{"issue":"22","key":"2024020116214416200_bib73","doi-asserted-by":"crossref","first-page":"2313","DOI":"10.1056\/NEJMoa0802885","article-title":"Stromal gene signatures in large-B-cell lymphomas","volume":"359","author":"Lenz","year":"2008","journal-title":"N\u00a0Engl J Med"},{"issue":"12","key":"2024020116214416200_bib74","doi-asserted-by":"crossref","first-page":"3151","DOI":"10.1084\/jem.20180668","article-title":"Loss of human ICOSL results in combined immunodeficiency","volume":"215","author":"Roussel","year":"2018","journal-title":"J\u00a0Exp Med"},{"issue":"1","key":"2024020116214416200_bib75","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1097\/00000478-200301000-00003","article-title":"Senile EBV+ B-cell lymphoproliferative disorders: a clinicopathologic study of 22 patients","volume":"27","author":"Oyama","year":"2003","journal-title":"Am J Surg Pathol"},{"issue":"17","key":"2024020116214416200_bib76","doi-asserted-by":"crossref","first-page":"5124","DOI":"10.1158\/1078-0432.CCR-06-2823","article-title":"Age-related EBV-associated B-cell lymphoproliferative disorders constitute a distinct clinicopathologic group: a study of 96\u00a0patients","volume":"13","author":"Oyama","year":"2007","journal-title":"Clin Cancer Res"},{"key":"2024020116214416200_bib77","doi-asserted-by":"crossref","first-page":"14340","DOI":"10.1038\/ncomms14340","article-title":"Intratumoral modulation of the inducible co-stimulator ICOS by recombinant oncolytic virus promotes systemic anti-tumour immunity","volume":"8","author":"Zamarin","year":"2017","journal-title":"Nat Commun"},{"issue":"11","key":"2024020116214416200_bib78","doi-asserted-by":"crossref","first-page":"1878","DOI":"10.1016\/j.ymthe.2019.07.013","article-title":"ICOS costimulation at the tumor site in combination with CTLA-4 blockade therapy elicits strong tumor immunity","volume":"27","author":"Soldevilla","year":"2019","journal-title":"Mol Ther"},{"issue":"7","key":"2024020116214416200_bib79","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1038\/nrd.2018.75","article-title":"The promise and challenges of immune agonist antibody development in cancer","volume":"17","author":"Mayes","year":"2018","journal-title":"Nat Rev Drug Discov"},{"issue":"17","key":"2024020116214416200_bib80","doi-asserted-by":"crossref","first-page":"3695","DOI":"10.1158\/1078-0432.CCR-21-4256","article-title":"First-in-human phase I\/II ICONIC Trial of the ICOS agonist vopratelimab alone and with nivolumab: ICOS-high CD4 T-cell populations and predictors of response","volume":"28","author":"Yap","year":"2022","journal-title":"Clin Cancer Res"},{"issue":"17","key":"2024020116214416200_bib81","doi-asserted-by":"crossref","first-page":"3633","DOI":"10.1158\/1078-0432.CCR-22-1520","article-title":"Agonizing over the stimulatory immune checkpoint ICOS","volume":"28","author":"Lee","year":"2022","journal-title":"Clin Cancer Res"}],"container-title":["Blood"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/ashpublications.org\/blood\/article-pdf\/143\/5\/429\/2210955\/blood_bld-2023-021346-main.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/ashpublications.org\/blood\/article-pdf\/143\/5\/429\/2210955\/blood_bld-2023-021346-main.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,2,1]],"date-time":"2024-02-01T16:24:03Z","timestamp":1706804643000},"score":1,"resource":{"primary":{"URL":"https:\/\/ashpublications.org\/blood\/article\/143\/5\/429\/498330\/Epstein-Barr-virus-encoded-EBNA2-downregulates"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,2,1]]},"references-count":82,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2024,2,1]]}},"URL":"https:\/\/doi.org\/10.1182\/blood.2023021346","relation":{},"ISSN":["0006-4971","1528-0020"],"issn-type":[{"value":"0006-4971","type":"print"},{"value":"1528-0020","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,2,1]]}}}