{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,31]],"date-time":"2025-10-31T07:57:58Z","timestamp":1761897478448,"version":"3.40.5"},"reference-count":86,"publisher":"Public Library of Science (PLoS)","issue":"2","license":[{"start":{"date-parts":[[2021,2,4]],"date-time":"2021-02-04T00:00:00Z","timestamp":1612396800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>Homeobox genes encode transcription factors which regulate basic processes in development and cell differentiation and are grouped into classes and subclasses according to sequence similarities. Here, we analyzed the activities of the 20 members strong TALE homeobox gene class in early hematopoiesis and in lymphopoiesis including developing and mature B-cells, T-cells, natural killer (NK)-cells and innate lymphoid cells (ILC). The resultant expression pattern comprised eleven genes and which we termed TALE-code enables discrimination of normal and aberrant activities of TALE homeobox genes in lymphoid malignancies. Subsequent expression analysis of TALE homeobox genes in public datasets of Hodgkin lymphoma (HL) patients revealed overexpression of IRX3, IRX4, MEIS1, MEIS3, PBX1, PBX4 and TGIF1. As paradigm we focused on PBX1 which was deregulated in about 17% HL patients. Normal PBX1 expression was restricted to hematopoietic stem cells and progenitors of T-cells and ILCs but absent in B-cells, reflecting its roles in stemness and early differentiation. HL cell line SUP-HD1 expressed enhanced PBX1 levels and served as an in vitro model to identify upstream regulators and downstream targets in this malignancy. Genomic studies of this cell line therein showed a gain of the PBX1 locus at 1q23 which may underlie its aberrant expression. Comparative expression profiling analyses of HL patients and cell lines followed by knockdown experiments revealed NFIB and TLX2 as target genes activated by PBX1. HOX proteins operate as cofactors of PBX1. Accordingly, our data showed that HOXB9 overexpressed in HL coactivated TLX2 but not NFIB while activating TNFRSF9 without PBX1. Further downstream analyses showed that TLX2 activated TBX15 which operated anti-apoptotically. Taken together, we discovered a lymphoid TALE-code and identified an aberrant network around deregulated TALE homeobox gene PBX1 which may disturb B-cell differentiation in HL by reactivation of progenitor-specific genes. These findings may provide the framework for future studies to exploit possible vulnerabilities of malignant cells in therapeutic scenarios.<\/jats:p>","DOI":"10.1371\/journal.pone.0246603","type":"journal-article","created":{"date-parts":[[2021,2,5]],"date-time":"2021-02-05T05:25:05Z","timestamp":1612502705000},"page":"e0246603","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":23,"title":["Establishment of the TALE-code reveals aberrantly activated homeobox gene PBX1 in Hodgkin lymphoma"],"prefix":"10.1371","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4431-8988","authenticated-orcid":true,"given":"Stefan","family":"Nagel","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Claudia","family":"Pommerenke","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Corinna","family":"Meyer","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Roderick A. F.","family":"MacLeod","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hans G.","family":"Drexler","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2021,2,4]]},"reference":[{"issue":"6","key":"pone.0246603.ref001","doi-asserted-by":"crossref","first-page":"2203","DOI":"10.1172\/JCI78083","article-title":"Checkpoints that control B cell development","volume":"125","author":"F Melchers","year":"2015","journal-title":"J Clin Invest"},{"issue":"1","key":"pone.0246603.ref002","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1111\/imr.12206","article-title":"The regulatory network of B-cell differentiation: a focused view of early B-cell factor 1 function","volume":"261","author":"S Boller","year":"2014","journal-title":"Immunol 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critical role in embryonic development of the axial skeleton and spleen","volume":"126","author":"C Tribioli","year":"1999","journal-title":"Development"},{"issue":"5","key":"pone.0246603.ref013","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1182\/blood-2014-06-579813","article-title":"A crucial role for the homeodomain transcription factor Hhex in lymphopoiesis","volume":"125","author":"JT Jackson","year":"2015","journal-title":"Blood"},{"issue":"4","key":"pone.0246603.ref014","doi-asserted-by":"crossref","first-page":"1531","DOI":"10.4049\/jimmunol.154.4.1531","article-title":"Perturbed development of T and B cells in mice expressing an Hlx homeobox transgene","volume":"154","author":"JD Allen","year":"1995","journal-title":"J Immunol"},{"key":"pone.0246603.ref015","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1186\/1741-7007-5-47","article-title":"Classification and nomenclature of all human homeobox genes","volume":"5","author":"PW Holland","year":"2007","journal-title":"BMC Biol"},{"issue":"21","key":"pone.0246603.ref016","doi-asserted-by":"crossref","first-page":"4173","DOI":"10.1093\/nar\/25.21.4173","article-title":"Analysis of TALE superclass homeobox genes (MEIS, PBC, KNOX, Iroquois, TGIF) reveals a novel domain conserved between plants and animals","volume":"25","author":"TR B\u00fcrglin","year":"1997","journal-title":"Nucleic Acids Res"},{"issue":"2","key":"pone.0246603.ref017","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1007\/s00239-006-0023-0","article-title":"Comprehensive analysis of animal TALE homeobox genes: new conserved motifs and cases of accelerated evolution","volume":"65","author":"K Mukherjee","year":"2007","journal-title":"J Mol Evol"},{"issue":"2","key":"pone.0246603.ref018","doi-asserted-by":"crossref","first-page":"e0171164","DOI":"10.1371\/journal.pone.0171164","article-title":"NKL homeobox gene activities in hematopoietic stem cells, T-cell development and T-cell 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Cancer"},{"issue":"4","key":"pone.0246603.ref022","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1182\/blood-2002-03-0839","article-title":"Loss of the B-lineage-specific gene expression program in Hodgkin and Reed-Sternberg cells of Hodgkin lymphoma","volume":"101","author":"I Schwering","year":"2003","journal-title":"Blood"},{"issue":"3","key":"pone.0246603.ref023","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/leu.2012.280","article-title":"Role of early B-cell factor 1 (EBF1) in Hodgkin lymphoma","volume":"27","author":"V Bohle","year":"2013","journal-title":"Leukemia"},{"issue":"2","key":"pone.0246603.ref024","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1038\/ni1285","article-title":"Intrinsic inhibition of transcription factor E2A by HLH proteins ABF-1 and Id2 mediates reprogramming of neoplastic B cells in Hodgkin lymphoma","volume":"7","author":"S Mathas","year":"2006","journal-title":"Nat 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Du","year":"2006","journal-title":"Genomics"},{"issue":"12","key":"pone.0246603.ref028","doi-asserted-by":"crossref","first-page":"1282","DOI":"10.1038\/ni.3299","article-title":"Long non-coding RNA profiling of human lymphoid progenitor cells reveals transcriptional divergence of B cell and T cell lineages","volume":"16","author":"D Casero","year":"2015","journal-title":"Nat Immunol"},{"issue":"6","key":"pone.0246603.ref029","doi-asserted-by":"crossref","first-page":"1086","DOI":"10.1016\/j.cell.2017.02.021","article-title":"Systemic human ILC precursors provide a substrate for tissue ILC differentiation","volume":"168","author":"AI Lim","year":"2017","journal-title":"Cell"},{"issue":"6","key":"pone.0246603.ref030","doi-asserted-by":"crossref","first-page":"2321","DOI":"10.1016\/j.jaci.2019.01.047","article-title":"Gene expression signatures of circulating human type 1, 2, and 3 innate lymphoid cells","volume":"143","author":"S Li","year":"2019","journal-title":"J Allergy Clin 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Nagel","year":"2018","journal-title":"Oncotarget"},{"issue":"1","key":"pone.0246603.ref034","doi-asserted-by":"crossref","first-page":"8218","DOI":"10.1038\/s41598-019-44491-x","article-title":"The LL-100 panel: 100 cell lines for blood cancer studies","volume":"9","author":"H Quentmeier","year":"2019","journal-title":"Sci Rep"},{"key":"pone.0246603.ref035","volume-title":"Guide to Leukemia-Lymphoma Cell Lines","author":"HG Drexler","year":"2010","edition":"2"},{"issue":"12","key":"pone.0246603.ref036","doi-asserted-by":"crossref","first-page":"1901","DOI":"10.1038\/sj.leu.2401592","article-title":"Standardized RT-PCR analysis of fusion gene transcripts from chromosome aberrations in acute leukemia for detection of minimal residual disease. Report of the BIOMED-1 Concerted Action: investigation of minimal residual disease in acute leukemia","volume":"13","author":"JJ van Dongen","year":"1999","journal-title":"Leukemia"},{"key":"pone.0246603.ref037","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1007\/978-1-4939-6703-2_5","article-title":"Cytogenetic harvesting of cancer cells and cell lines","volume":"1541","author":"RA MacLeod","year":"2017","journal-title":"Methods Mol Biol"},{"issue":"5","key":"pone.0246603.ref038","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1016\/j.stem.2008.03.004","article-title":"Pbx1 regulates self-renewal of long-term hematopoietic stem cells by maintaining their quiescence","volume":"2","author":"F Ficara","year":"2008","journal-title":"Cell Stem Cell"},{"issue":"Pt 14","key":"pone.0246603.ref039","doi-asserted-by":"crossref","first-page":"3181","DOI":"10.1242\/jcs.125435","article-title":"Pbx1 restrains myeloid maturation while preserving lymphoid potential in hematopoietic progenitors","volume":"126","author":"F Ficara","year":"2013","journal-title":"J Cell Sci"},{"issue":"10","key":"pone.0246603.ref040","doi-asserted-by":"crossref","first-page":"4191","DOI":"10.1182\/blood-2006-10-054213","article-title":"B-cell development fails in the absence of the Pbx1 proto-oncogene","volume":"109","author":"M Sanyal","year":"2007","journal-title":"Blood"},{"issue":"3","key":"pone.0246603.ref041","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1007\/s00018-017-2620-z","article-title":"Molecular processes involved in B cell acute lymphoblastic leukaemia","volume":"75","author":"C Malouf","year":"2018","journal-title":"Cell Mol Life Sci"},{"issue":"4","key":"pone.0246603.ref042","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1016\/0092-8674(90)90658-2","article-title":"A new homeobox gene contributes the DNA binding domain of the t(1;19) translocation protein in pre-B ALL","volume":"60","author":"MP Kamps","year":"1990","journal-title":"Cell"},{"key":"pone.0246603.ref043","first-page":"25","article-title":"E2A-Pbx1 induces growth, blocks differentiation, and interacts with other homeodomain proteins regulating normal differentiation","volume":"220","author":"MP Kamps","year":"1997","journal-title":"Curr Top Microbiol Immunol"},{"key":"pone.0246603.ref044","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.canlet.2017.09.015","article-title":"The convergent roles of the nuclear factor I transcription factors in development and cancer","volume":"410","author":"KS Chen","year":"2017","journal-title":"Cancer Lett"},{"issue":"13","key":"pone.0246603.ref045","doi-asserted-by":"crossref","first-page":"3113","DOI":"10.1242\/dev.01884","article-title":"A Pbx1-dependent genetic and transcriptional network regulates spleen ontogeny","volume":"132","author":"A Brendolan","year":"2005","journal-title":"Development"},{"issue":"5","key":"pone.0246603.ref046","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1007\/s004290050061","article-title":"Ncx, a Hox11 related gene, is expressed in a variety of tissues derived from neural crest cells","volume":"195","author":"M Hatano","year":"1997","journal-title":"Anat Embryol"},{"issue":"4","key":"pone.0246603.ref047","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1182\/blood.V97.4.1063","article-title":"Expression of the c-met proto-oncogene and its ligand, hepatocyte growth factor, in Hodgkin disease","volume":"97","author":"L Teofili","year":"2001","journal-title":"Blood"},{"key":"pone.0246603.ref048","doi-asserted-by":"crossref","first-page":"7596","DOI":"10.1038\/srep07596","article-title":"Smad9 is a new type of transcriptional regulator in bone morphogenetic protein signaling","volume":"4","author":"S Tsukamoto","year":"2014","journal-title":"Sci Rep"},{"issue":"3","key":"pone.0246603.ref049","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1016\/j.molcel.2012.01.002","article-title":"PCGF homologs, CBX proteins, and RYBP define functionally distinct PRC1 family complexes","volume":"45","author":"Z Gao","year":"2012","journal-title":"Mol Cell"},{"issue":"8","key":"pone.0246603.ref050","doi-asserted-by":"crossref","first-page":"537","DOI":"10.4161\/epi.4.8.10132","article-title":"Epigenetic regulation of vertebrate Hox genes: a dynamic equilibrium","volume":"4","author":"N Soshnikova","year":"2009","journal-title":"Epigenetics"},{"key":"pone.0246603.ref051","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1016\/bs.ctdb.2016.08.009","article-title":"The roles of T-box genes in vertebrate limb development","volume":"122","author":"CJ Sheeba","year":"2017","journal-title":"Curr Top Dev Biol"},{"issue":"10","key":"pone.0246603.ref052","doi-asserted-by":"crossref","first-page":"1338","DOI":"10.1007\/s10495-015-1155-8","article-title":"Novel antiapoptotic effect of TBX15: overexpression of TBX15 reduces apoptosis in cancer cells","volume":"20","author":"J Arribas","year":"2015","journal-title":"Apoptosis"},{"issue":"2","key":"pone.0246603.ref053","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.ydbio.2005.10.032","article-title":"Hox cofactors in vertebrate development","volume":"291","author":"CB Moens","year":"2006","journal-title":"Dev Biol"},{"issue":"4","key":"pone.0246603.ref054","doi-asserted-by":"crossref","first-page":"1734","DOI":"10.1128\/MCB.16.4.1734","article-title":"Pbx modulation of Hox homeodomain amino-terminal arms establishes different DNA-binding specificities across the Hox locus","volume":"16","author":"CP Chang","year":"1996","journal-title":"Mol Cell Biol"},{"issue":"7","key":"pone.0246603.ref055","doi-asserted-by":"crossref","first-page":"3015","DOI":"10.1182\/blood-2006-08-044347","article-title":"Comprehensive analysis of homeobox genes in Hodgkin lymphoma cell lines identifies dysregulated expression of HOXB9 mediated via ERK5 signaling and BMI1","volume":"109","author":"S Nagel","year":"2007","journal-title":"Blood"},{"issue":"2","key":"pone.0246603.ref056","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1158\/0008-5472.CAN-12-3849","article-title":"Expression of CD137 on Hodgkin and Reed-Sternberg cells inhibits T-cell activation by eliminating CD137 ligand expression","volume":"73","author":"WT Ho","year":"2013","journal-title":"Cancer Res"},{"key":"pone.0246603.ref057","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.ebiom.2018.11.053","article-title":"Mapping of \u03b3\/\u03b4 T cells reveals V\u03b42+ T cells resistance to senescence","volume":"39","author":"W Xu","year":"2019","journal-title":"EBioMedicine"},{"issue":"5","key":"pone.0246603.ref058","doi-asserted-by":"crossref","first-page":"6479","DOI":"10.1096\/fj.202000121R","article-title":"PBX1 promotes development of natural killer cells by binding directly to the Nfil3 promoter","volume":"34","author":"X Xu","year":"2020","journal-title":"FASEB J"},{"issue":"7","key":"pone.0246603.ref059","doi-asserted-by":"crossref","first-page":"558","DOI":"10.1002\/gcc.20559","article-title":"Detection of a t(1;22)(q23;q12) translocation leading to an EWSR1-PBX1 fusion gene in a myoepithelioma","volume":"47","author":"P Brandal","year":"2008","journal-title":"Genes Chromosomes Cancer"},{"issue":"5","key":"pone.0246603.ref060","doi-asserted-by":"crossref","first-page":"e36054","DOI":"10.1371\/journal.pone.0036054","article-title":"Identification of PBX1 target genes in cancer cells by global mapping of PBX1 binding sites","volume":"7","author":"MM Thiaville","year":"2012","journal-title":"PLoS One"},{"key":"pone.0246603.ref061","doi-asserted-by":"crossref","first-page":"9","DOI":"10.3389\/fcell.2017.00009","article-title":"PBX1 as Pioneer Factor: A case still open","volume":"5","author":"BM Grebbin","year":"2017","journal-title":"Front Cell Dev Biol"},{"issue":"18","key":"pone.0246603.ref062","doi-asserted-by":"crossref","first-page":"3543","DOI":"10.1242\/dev.128.18.3543","article-title":"Requirement for Pbx1 in skeletal patterning and programming chondrocyte proliferation and differentiation","volume":"128","author":"L Selleri","year":"2001","journal-title":"Development"},{"issue":"3","key":"pone.0246603.ref063","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1111\/j.1600-0609.1988.tb00823.x","article-title":"Partial duplication of chromosome 1q preceding the development of an L3 lymphoblastic leukemia with t(8;14), secondary to treatment for Hodgkin\u2019s disease","volume":"40","author":"M Zollino","year":"1988","journal-title":"Eur J Haematol"},{"issue":"4","key":"pone.0246603.ref064","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1002\/gcc.2870030408","article-title":"Clonal cytogenetic abnormalities in Hodgkin\u2019s disease","volume":"3","author":"M Ladanyi","year":"1991","journal-title":"Genes Chromosomes Cancer"},{"issue":"4","key":"pone.0246603.ref065","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1002\/gcc.2870050415","article-title":"Recurring chromosome abnormalities in Hodgkin\u2019s disease","volume":"5","author":"H D\u00f6hner","year":"1992","journal-title":"Genes Chromosomes Cancer"},{"issue":"11","key":"pone.0246603.ref066","doi-asserted-by":"crossref","first-page":"e40","DOI":"10.1038\/bcj.2011.39","article-title":"Analysis of genomic aberrations and gene expression profiling identifies novel lesions and pathways in myeloproliferative neoplasms","volume":"1","author":"KL Rice","year":"2011","journal-title":"Blood Cancer J"},{"key":"pone.0246603.ref067","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.ebiom.2017.05.027","article-title":"Nuclear Factor I\/B: A Master regulator of cell differentiation with paradoxical roles in cancer","volume":"22","author":"DD Becker-Santos","year":"2017","journal-title":"EBioMedicine"},{"issue":"6204","key":"pone.0246603.ref068","doi-asserted-by":"crossref","first-page":"1251033","DOI":"10.1126\/science.1251033","article-title":"Transcriptional diversity during lineage commitment of human blood progenitors","volume":"345","author":"L Chen","year":"2014","journal-title":"Science"},{"issue":"17","key":"pone.0246603.ref069","doi-asserted-by":"crossref","first-page":"2987","DOI":"10.1182\/blood-2013-04-493973","article-title":"Nfix is a novel regulator of murine hematopoietic stem and progenitor cell survival","volume":"122","author":"P Holmfeldt","year":"2013","journal-title":"Blood"},{"issue":"9","key":"pone.0246603.ref070","doi-asserted-by":"crossref","first-page":"1753","DOI":"10.1182\/blood-2008-12-196196","article-title":"NFI-A directs the fate of hematopoietic progenitors to the erythroid or granulocytic lineage and controls beta-globin and G-CSF receptor expression","volume":"114","author":"LM Starnes","year":"2009","journal-title":"Blood"},{"issue":"2","key":"pone.0246603.ref071","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1042\/BJ20051386","article-title":"The TLX2 homeobox gene is a transcriptional target of PHOX2B in neural-crest-derived cells","volume":"395","author":"S Borghini","year":"2006","journal-title":"Biochem J"},{"issue":"35","key":"pone.0246603.ref072","doi-asserted-by":"crossref","first-page":"3227","DOI":"10.18632\/oncotarget.26929","article-title":"Deregulated expression of NKL homeobox genes in T-cell lymphomas","volume":"10","author":"S Nagel","year":"2019","journal-title":"Oncotarget"},{"issue":"2","key":"pone.0246603.ref073","doi-asserted-by":"crossref","first-page":"480","DOI":"10.3109\/10428194.2014.924119","article-title":"Repressed BMP signaling reactivates NKL homeobox gene MSX1 in a T-ALL subset","volume":"56","author":"S Nagel","year":"2015","journal-title":"Leuk Lymphoma"},{"issue":"5","key":"pone.0246603.ref074","doi-asserted-by":"crossref","first-page":"e0197194","DOI":"10.1371\/journal.pone.0197194","article-title":"Aberrant activity of NKL homeobox gene NKX3-2 in a T-ALL subset","volume":"13","author":"S Nagel","year":"2018","journal-title":"PLoS One"},{"issue":"40","key":"pone.0246603.ref075","doi-asserted-by":"crossref","first-page":"25628","DOI":"10.1074\/jbc.273.40.25628","article-title":"Specific activation of Smad1 signaling pathways by the BMP7 type I receptor, ALK2","volume":"273","author":"M Mac\u00edas-Silva","year":"1998","journal-title":"J Biol Chem"},{"issue":"10","key":"pone.0246603.ref076","doi-asserted-by":"crossref","first-page":"1877","DOI":"10.1242\/dev.125.10.1877","article-title":"The Tlx-2 homeobox gene is a downstream target of BMP signalling and is required for mouse mesoderm development","volume":"125","author":"SJ Tang","year":"1998","journal-title":"Development"},{"issue":"17","key":"pone.0246603.ref077","first-page":"5329","article-title":"The cardiac homeobox gene NKX2-5 is deregulated by juxtaposition with BCL11B in pediatric T-ALL cell lines via a novel t(5;14)(q35.1;q32.2)","volume":"63","author":"S Nagel","year":"2003","journal-title":"Cancer Res"},{"issue":"8","key":"pone.0246603.ref078","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1038\/sj.leu.2400718","article-title":"Deletion of chromosome arm 3p in hematologic malignancies","volume":"11","author":"B Johansson","year":"1997","journal-title":"Leukemia"},{"issue":"5","key":"pone.0246603.ref079","doi-asserted-by":"crossref","first-page":"528","DOI":"10.1002\/path.1661","article-title":"Abnormal PcG protein expression in Hodgkin\u2019s lymphoma. Relation with E2F6 and NFkappaB transcription factors","volume":"204","author":"M S\u00e1nchez-Beato","year":"2004","journal-title":"J Pathol"},{"issue":"6","key":"pone.0246603.ref080","doi-asserted-by":"crossref","first-page":"e0157761","DOI":"10.1371\/journal.pone.0157761","article-title":"NF-kappaB mediates the expression of TBX15 in cancer cells","volume":"11","author":"J Arribas","year":"2016","journal-title":"PLoS One"},{"issue":"5\u20136","key":"pone.0246603.ref081","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1101\/gad.318774.118","article-title":"\u2019Building a perfect body\u2019: control of vertebrate organogenesis by PBX-dependent regulatory networks","volume":"33","author":"L Selleri","year":"2019","journal-title":"Genes Dev"},{"issue":"13","key":"pone.0246603.ref082","doi-asserted-by":"crossref","first-page":"8198","DOI":"10.1074\/jbc.272.13.8198","article-title":"The Abd-B-like Hox homeodomain proteins can be subdivided by the ability to form complexes with Pbx1a on a novel DNA target","volume":"272","author":"WF Shen","year":"1997","journal-title":"J Biol Chem"},{"issue":"11","key":"pone.0246603.ref083","doi-asserted-by":"crossref","first-page":"E3299","DOI":"10.3390\/cancers12113299","article-title":"HOX genes family and cancer: a novel role for Homeobox B9 in the resistance to anti-angiogenic therapies","volume":"12","author":"S Contarelli","year":"2020","journal-title":"Cancers (Basel)"},{"issue":"6","key":"pone.0246603.ref084","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1016\/j.tig.2016.03.004","article-title":"To Be Specific or Not: The critical relationship between Hox and TALE proteins","volume":"32","author":"S Merabet","year":"2016","journal-title":"Trends Genet"},{"issue":"1","key":"pone.0246603.ref085","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1038\/leu.2013.211","article-title":"PAX5 overexpression is not enough to reestablish the mature B-cell phenotype in classical Hodgkin lymphoma","volume":"28","author":"L Dimitrova","year":"2014","journal-title":"Leukemia"},{"issue":"19","key":"pone.0246603.ref086","doi-asserted-by":"crossref","first-page":"32322","DOI":"10.18632\/oncotarget.15971","article-title":"Targeting HOX\/PBX dimers in cancer","volume":"8","author":"R Morgan","year":"2017","journal-title":"Oncotarget"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0246603","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,10,20]],"date-time":"2023-10-20T09:37:23Z","timestamp":1697794643000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0246603"}},"subtitle":[],"editor":[{"given":"Annalisa","family":"Di Ruscio","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2021,2,4]]},"references-count":86,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2021,2,4]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0246603","relation":{},"ISSN":["1932-6203"],"issn-type":[{"type":"electronic","value":"1932-6203"}],"subject":[],"published":{"date-parts":[[2021,2,4]]}}}