{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T17:53:52Z","timestamp":1790704432205,"version":"4.1.0"},"reference-count":68,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2019,4,16]],"date-time":"2019-04-16T00:00:00Z","timestamp":1555372800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"DOI":"10.3389\/fimmu.2019.00568","type":"journal-article","created":{"date-parts":[[2019,4,16]],"date-time":"2019-04-16T01:29:21Z","timestamp":1555378161000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":54,"title":["Assessing the Functional Relevance of Variants in the IKAROS Family Zinc Finger Protein 1 (IKZF1) in a Cohort of Patients With Primary Immunodeficiency"],"prefix":"10.3389","volume":"10","author":[{"given":"Zoya","family":"Eskandarian","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manfred","family":"Fliegauf","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alla","family":"Bulashevska","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michele","family":"Proietti","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosie","family":"Hague","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cristian Roberto","family":"Smulski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Desir\u00e9e","family":"Schubert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Klaus","family":"Warnatz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bodo","family":"Grimbacher","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2019,4,16]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1182\/asheducation-2012.1.301","article-title":"The many faces of common variable immunodeficiency","volume":"2012","author":"Cunningham-Rundles","year":"2012","journal-title":"Hematology Am Soc Hematol Educ Program."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1650","DOI":"10.1182\/blood-2011-09-377945","article-title":"Morbidity and mortality in common variable immune deficiency over 4 decades","volume":"119","author":"Resnick","year":"2012","journal-title":"Blood."},{"key":"B3","doi-asserted-by":"publisher","first-page":"1544","DOI":"10.1182\/blood.V99.5.1544","article-title":"Severe deficiency of switched memory B cells (CD27+IgM\u2212IgD\u2212) in subgroups of patients with common variable immunodeficiency: a new approach to classify a heterogeneous disease","volume":"99","author":"Warnatz","year":"2002","journal-title":"Blood"},{"key":"B4","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1016\/j.jaci.2013.12.1077","article-title":"Clinical picture and treatment of 2212 patients with common variable immunodeficiency","volume":"134","author":"Gathmann","year":"2014","journal-title":"J Allergy Clin Immunol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1016\/j.jaip.2015.07.025","article-title":"International Consensus Document (ICON): common variable immunodeficiency disorders","volume":"4","author":"Bonilla","year":"2016","journal-title":"J Allergy Clin Immunol Pract."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1136\/bcr-2017-221657","article-title":"Patients with common variable immunodeficiency paradoxically have increased rates of autoimmune disorders","author":"Susheela","year":"2017","journal-title":"BMJ Case Rep."},{"key":"B7","doi-asserted-by":"publisher","first-page":"210","DOI":"10.3389\/fimmu.2012.00210","article-title":"Pathogenesis of autoimmunity in common variable immunodeficiency","volume":"3","author":"Warnatz","year":"2012","journal-title":"Front Immunol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1016\/j.leukres.2015.02.002","article-title":"Lymphoproliferative disease and cancer among patients with common variable immunodeficiency","volume":"39","author":"Gangemi","year":"2015","journal-title":"Leuk Res."},{"key":"B9","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1007\/s11882-013-0350-3","article-title":"Autoimmunity in Immunodeficiency","volume":"13","author":"Todoric","year":"2013","journal-title":"Curr Allergy Asthma Rep."},{"key":"B10","doi-asserted-by":"publisher","first-page":"220","DOI":"10.3389\/fimmu.2016.00220","article-title":"Genetic diagnosis using whole exome sequencing in common variable immunodeficiency","volume":"7","author":"Maffucci","year":"2016","journal-title":"Front Immunol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"812","DOI":"10.1016\/j.ajhg.2013.09.009","article-title":"Germline mutations in NFKB2 implicate the noncanonical NF-\u03baB pathway in the pathogenesis of common variable immunodeficiency","volume":"93","author":"Chen","year":"2013","journal-title":"Am J Hum Genet."},{"key":"B12","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/j.coi.2014.05.004","article-title":"Discovery of single-gene inborn errors of immunity by next generation sequencing","volume":"30","author":"Conley","year":"2014","journal-title":"Curr Opin Immunol."},{"key":"B13","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1016\/j.ajhg.2015.07.008","article-title":"Haploinsufficiency of the NF-\u03baB1 Subunit p50 in common variable immunodeficiency","volume":"97","author":"Fliegauf","year":"2015","journal-title":"Am J Hum Genet."},{"key":"B14","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1038\/ni.2771","article-title":"Dominant-activating, germline mutations in phosphoinositide 3-Kinase p110\u03b4 cause T cell senescence and human immunodeficiency","volume":"15","author":"Lucas","year":"2014","journal-title":"Nat Immunol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1007\/s10875-017-0464-9","article-title":"International union of immunological societies: 2017 primary immunodeficiency diseases committee report on inborn errors of immunity","volume":"38","author":"Picard","year":"2018","journal-title":"J Clin Immunol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.1056\/NEJMoa1512234","article-title":"Loss of B cells in patients with heterozygous mutations in IKAROS","volume":"374","author":"Kuehn","year":"2016","journal-title":"N Engl J Med."},{"key":"B17","doi-asserted-by":"publisher","first-page":"1272","DOI":"10.1016\/j.molimm.2011.03.006","article-title":"The Ikaros gene family: transcriptional regulators of hematopoiesis and immunity","volume":"48","author":"John","year":"2011","journal-title":"Mol Immunol"},{"key":"B18","doi-asserted-by":"publisher","first-page":"537","DOI":"10.1016\/S1074-7613(00)80269-1","article-title":"Selective defects in the development of the fetal and adult lymphoid system in mice with an ikaros null mutation","volume":"5","author":"Wang","year":"1996","journal-title":"Immunity."},{"key":"B19","doi-asserted-by":"publisher","first-page":"2146","DOI":"10.1101\/gad.816400","article-title":"Targeting of Ikaros to pericentromeric heterochromatin by direct DNA binding","volume":"14","author":"Cobb","year":"2000","journal-title":"Genes Dev"},{"key":"B20","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1016\/S1074-7613(03)00168-7","article-title":"Widespread failure of hematolymphoid differentiation caused by a recessive niche-filling allele of the ikaros transcription factor","volume":"19","author":"Papathanasiou","year":"2003","journal-title":"Immunity."},{"key":"B21","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/0092-8674(94)90407-3","article-title":"The ikaros gene is required for the development of all lymphoid lineages","volume":"79","author":"Georgopoulos","year":"1994","journal-title":"Cell."},{"key":"B22","doi-asserted-by":"publisher","first-page":"808","DOI":"10.1126\/science.1439790","article-title":"Ikaros, an early lymphoid-specific transcription factor and a putative mediator for T cell commitment","volume":"258","author":"Georgopoulos","year":"1992","journal-title":"Science."},{"key":"B23","doi-asserted-by":"publisher","first-page":"720","DOI":"10.1002\/1521-4141(200203)32:3<720::AID-IMMU720>3.0.CO;2-P","article-title":"Ikaros is critical for B cell differentiation and function","volume":"32","author":"Kirstetter","year":"2002","journal-title":"Eur J Immunol."},{"key":"B24","doi-asserted-by":"publisher","first-page":"1239","DOI":"10.1038\/s41467-018-02977-8","article-title":"Ikaros family zinc finger 1 regulates dendritic cell development and function in humans","volume":"9","author":"Cytlak","year":"2018","journal-title":"Nat Commun."},{"key":"B25","doi-asserted-by":"publisher","first-page":"2211","DOI":"10.1093\/emboj\/cdg186","article-title":"A complex network of regulatory elements in Ikaros and their activity during hemo-lymphopoiesis","volume":"22","author":"Kaufmann","year":"2003","journal-title":"EMBO J."},{"key":"B26","doi-asserted-by":"publisher","first-page":"826","DOI":"10.1046\/j.1365-2141.2000.02297.x","article-title":"Detection of different Ikaros isoforms in human leukaemias using real-time quantitative polymerase chain reaction","volume":"110","author":"Olivero","year":"2000","journal-title":"Br J Haematol."},{"key":"B27","doi-asserted-by":"publisher","first-page":"773","DOI":"10.1038\/nri3550","article-title":"Two versions of the Ikaros tale: haematopoiesis","volume":"13","author":"Papatriantafyllou","year":"2013","journal-title":"Nat Rev Immunol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"161","DOI":"10.4331\/wjbc.v2.i6.161","article-title":"Zinc finger structure-function in Ikaros","volume":"2","author":"Payne","year":"2011","journal-title":"World J Biol Chem."},{"key":"B29","doi-asserted-by":"publisher","first-page":"1073","DOI":"10.1038\/ni.2707","article-title":"Selective regulation of lymphopoiesis and leukemogenesis by individual zinc fingers of Ikaros","volume":"14","author":"Schjerven","year":"2013","journal-title":"Nat Immunol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"1817","DOI":"10.1101\/gad.905601","article-title":"Down-regulation of TDT transcription in CD4+CD8+ thymocytes by Ikaros proteins in direct competition with an Ets activator","volume":"15","author":"Trinh","year":"2001","journal-title":"Genes Dev"},{"key":"B31","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1128\/MCB.26.1.209-220.2006","article-title":"Notch activation is an early and critical event during T-cell leukemogenesis in ikaros-deficient mice","volume":"26","author":"Dumortier","year":"2006","journal-title":"Mol Cell Biol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"1201","DOI":"10.1084\/jem.190.9.1201","article-title":"Defects in hemopoietic stem cell activity in ikaros mutant mice","volume":"190","author":"Nichogiannopoulou","year":"1999","journal-title":"J Exp Med."},{"key":"B33","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1002\/pbc.23160","article-title":"Congenital pancytopenia and absence of B lymphocytes in a neonate with a mutation in the ikaros gene","volume":"58","author":"Goldman","year":"2012","journal-title":"Pediatr Blood Cancer."},{"key":"B34","doi-asserted-by":"publisher","first-page":"432","DOI":"10.1016\/j.jaci.2017.08.019","article-title":"A novel IKAROS haploinsufficiency kindred with unexpectedly late and variable B-cell maturation defects","volume":"141","author":"Bogaert","year":"2018","journal-title":"J Allergy Clin Immunol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1016\/j.jaci.2016.09.029","article-title":"Abnormal hematopoiesis and autoimmunity in human subjects with germline IKZF1 mutations","volume":"140","author":"Hoshino","year":"2017","journal-title":"J Allergy Clin Immunol."},{"key":"B36","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1016\/j.jaci.2018.04.008","article-title":"A kindred with mutant IKAROS and autoimmunity","volume":"142","author":"Van Nieuwenhove","year":"2018","journal-title":"J Allergy Clin Immunol."},{"key":"B37","doi-asserted-by":"publisher","first-page":"1221","DOI":"10.1038\/leu.2017.25","article-title":"Germline IKAROS mutation associated with primary immunodeficiency that progressed to T-cell acute lymphoblastic leukemia","volume":"31","author":"Yoshida","year":"2017","journal-title":"Leukemia."},{"key":"B38","doi-asserted-by":"publisher","first-page":"3071","DOI":"10.1172\/JCI98164","article-title":"Dominant-negative IKZF1 mutations cause a T, B, and myeloid cell combined immunodeficiency","volume":"128","author":"Boutboul","year":"2018","journal-title":"J Clin Invest."},{"key":"B39","doi-asserted-by":"publisher","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":"B40","doi-asserted-by":"publisher","first-page":"1297","DOI":"10.1101\/gr.107524.110","article-title":"The genome analysis toolkit: a MapReduce framework for analyzing next-generation DNA sequencing data","volume":"20","author":"McKenna","year":"2010","journal-title":"Genome Res"},{"key":"B41","doi-asserted-by":"publisher","first-page":"2156","DOI":"10.1093\/bioinformatics\/btr330","article-title":"The variant call format and VCFtools","volume":"27","author":"Danecek","year":"2011","journal-title":"Bioinformatics."},{"key":"B42","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1038\/sj.gene.6364173","article-title":"Immune response in silico (IRIS): immune-specific genes identified from a compendium of microarray expression data","volume":"6","author":"Abbas","year":"2005","journal-title":"Genes Immun."},{"key":"B43","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1038\/75556","article-title":"Gene ontology: tool for the unification of biology","volume":"25","author":"Ashburner","year":"2000","journal-title":"Nat Genet."},{"key":"B44","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/B978-0-12-801430-1.00005-6","article-title":"Tools and techniques to study ligand-receptor interactions and receptor activation by TNF superfamily members","volume":"545","author":"Schneider","year":"2014","journal-title":"Meth Enzymol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"e2601","DOI":"10.1038\/cddis.2017.22","article-title":"Hetero-oligomerization between the TNF receptor superfamily members CD40, Fas and TRAILR2 modulate CD40 signalling","volume":"8","author":"Smulski","year":"2017","journal-title":"Cell Death Dis."},{"key":"B46","doi-asserted-by":"publisher","first-page":"8292","DOI":"10.1128\/MCB.14.12.8292","article-title":"The Ikaros gene encodes a family of functionally diverse zinc finger DNA-binding proteins","volume":"14","author":"Moln\u00e1r","year":"1994","journal-title":"Mol Cell Biol."},{"key":"B47","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1016\/S1097-2765(03)00043-1","article-title":"Selective dimerization of a C2H2 zinc finger subfamily","volume":"11","author":"McCarty","year":"2003","journal-title":"Mol Cell."},{"key":"B48","doi-asserted-by":"publisher","first-page":"2985","DOI":"10.1101\/gad.1040502","article-title":"A common mechanism for mitotic inactivation of C2H2 zinc finger DNA-binding domains","volume":"16","author":"Dovat","year":"2002","journal-title":"Genes Dev."},{"key":"B49","doi-asserted-by":"publisher","first-page":"8291","DOI":"10.1074\/jbc.M707906200","article-title":"Recruitment of ikaros to pericentromeric heterochromatin is regulated by phosphorylation","volume":"283","author":"Gurel","year":"2008","journal-title":"J Biol Chem"},{"key":"B50","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1101\/gr.086496.108","article-title":"Human gamma-satellite DNA maintains open chromatin structure and protects a transgene from epigenetic silencing","volume":"19","author":"Kim","year":"2009","journal-title":"Genome Res."},{"key":"B51","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1007\/BF00661276","article-title":"Isolation and identification of a novel tandemly repeated DNA sequence in the centromeric region of human chromosome 8","volume":"102","author":"Lin","year":"1993","journal-title":"Chromosoma."},{"key":"B52","doi-asserted-by":"publisher","first-page":"10563","DOI":"10.1073\/pnas.0503346102","article-title":"Progressive proximal expansion of the primate X chromosome centromere","volume":"102","author":"Schueler","year":"2005","journal-title":"Proc Natl Acad Sci USA."},{"key":"B53","doi-asserted-by":"publisher","first-page":"5118","DOI":"10.4049\/jimmunol.1301992","article-title":"Ikaros imposes a barrier to CD8+ T cell differentiation by restricting autocrine IL-2 production","volume":"192","author":"O'Brien","year":"2014","journal-title":"J Immunol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"e40934","DOI":"10.1371\/journal.pone.0040934","article-title":"IKAROS deletions dictate a unique gene expression signature in patients with adult B-cell acute lymphoblastic leukemia","volume":"7","author":"Iacobucci","year":"","journal-title":"PLoS ONE."},{"key":"B55","first-page":"1","article-title":"Function of Ikaros as a tumor suppressor in B cell acute lymphoblastic leukemia","volume":"3","author":"Kastner","year":"2013","journal-title":"Am J Blood Res."},{"key":"B56","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1056\/NEJMoa0808253","article-title":"Deletion of IKZF1 and prognosis in acute lymphoblastic leukemia","volume":"360","author":"Mullighan","year":"2009","journal-title":"N Engl J Med."},{"key":"B57","doi-asserted-by":"publisher","first-page":"680","DOI":"10.1073\/pnas.96.2.680","article-title":"Expression of dominant-negative and mutant isoforms of the antileukemic transcription factor Ikaros in infant acute lymphoblastic leukemia","volume":"96","author":"Sun","year":"1999","journal-title":"Proc Natl Acad Sci USA."},{"key":"B58","doi-asserted-by":"publisher","first-page":"1560","DOI":"10.1126\/science.1260791","article-title":"Autoimmunity by haploinsufficiency","volume":"345","author":"Rieux-Laucat","year":"2014","journal-title":"Science."},{"key":"B59","doi-asserted-by":"publisher","first-page":"937","DOI":"10.1189\/jlb.0510246","article-title":"B cell receptor triggering sensitizes human B cells to TRAIL-induced apoptosis","volume":"88","author":"Guerreiro-Cacais","year":"2010","journal-title":"J Leuk Biol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"5505","DOI":"10.4049\/jimmunol.169.10.5505","article-title":"TRAIL\/Apo-2 ligand induces primary plasma cell apoptosis","volume":"169","author":"Ursini-Siegel","year":"2002","journal-title":"J Immunol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"1091","DOI":"10.1517\/14728222.2010.519701","article-title":"TRAIL receptor signaling and therapeutics","volume":"14","author":"Abdulghani","year":"2010","journal-title":"Expert Opin Ther Targets."},{"key":"B62","doi-asserted-by":"publisher","first-page":"673","DOI":"10.1016\/1074-7613(95)90057-8","article-title":"Identification and characterization of a new member of the TNF family that induces apoptosis","volume":"3","author":"Wiley","year":"1995","journal-title":"Immunity."},{"key":"B63","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/S1043-4666(03)00094-2","article-title":"Regulation of soluble and surface-bound TRAIL in human T cells, B cells, and monocytes","volume":"24","author":"Ehrlich","year":"2003","journal-title":"Cytokine."},{"key":"B64","doi-asserted-by":"publisher","first-page":"2418","DOI":"10.1182\/blood-2004-04-1294","article-title":"Activated human NK and CD8+ T cells express both TNF-related apoptosis-inducing ligand (TRAIL) and TRAIL receptors but are resistant to TRAIL-mediated cytotoxicity","volume":"104","author":"Mirandola","year":"2004","journal-title":"Blood."},{"key":"B65","doi-asserted-by":"publisher","first-page":"1462","DOI":"10.1016\/j.biocel.2007.02.007","article-title":"TRAIL signalling: Decisions between life and death","volume":"39","author":"Falschlehner","year":"2007","journal-title":"Int J Biochem Cell Biol."},{"key":"B66","doi-asserted-by":"publisher","first-page":"831","DOI":"10.1016\/S1074-7613(00)80401-X","article-title":"TRAIL receptors 1 (DR4) and 2 (DR5) signal FADD-dependent apoptosis and activate NF-\u03baB","volume":"7","author":"Schneider","year":"1997","journal-title":"Immunity."},{"key":"B67","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1016\/j.autrev.2012.10.017","article-title":"Chipping away at a mountain: genomic studies in common variable immunodeficiency","volume":"12","author":"Keller","year":"2013","journal-title":"Autoimmun Rev."},{"key":"B68","doi-asserted-by":"publisher","first-page":"1360","DOI":"10.1016\/j.jaci.2011.02.039","article-title":"Genome-wide association identifies diverse causes of common variable immunodeficiency","volume":"127","author":"Orange","year":"2011","journal-title":"J Allergy Clin Immunol."}],"updated-by":[{"DOI":"10.3389\/fimmu.2019.01490","type":"corrigendum","label":"Corrigendum","source":"publisher","updated":{"date-parts":[[2019,6,28]],"date-time":"2019-06-28T00:00:00Z","timestamp":1561680000000}}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fimmu.2019.00568\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2019,6,28]],"date-time":"2019-06-28T10:05:04Z","timestamp":1561716304000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fimmu.2019.00568\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,4,16]]},"references-count":68,"alternative-id":["10.3389\/fimmu.2019.00568"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2019.00568","relation":{"corrigendum":[{"id-type":"doi","id":"10.3389\/fimmu.2019.01490","asserted-by":"object"}]},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,4,16]]},"article-number":"568"}}