{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T09:57:46Z","timestamp":1790675866985,"version":"4.1.0"},"reference-count":77,"publisher":"Wiley","issue":"9","license":[{"start":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T00:00:00Z","timestamp":1789084800000},"content-version":"vor","delay-in-days":10,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["19H03623"],"award-info":[{"award-number":["19H03623"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["BioEssays"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Neural progenitor cells (NPCs) must preserve lineage identity while remaining responsive to developmental cues. Here, we discuss the hypothesis that ATRX condensates help organize enhancer\u2010centered nuclear microenvironments in NPCs. ATRX has long been studied in heterochromatin maintenance, histone variant deposition, and chromatin remodeling; earlier work has also shown that ATRX can occupy euchromatic and active regulatory regions and contribute to transcriptional regulation. Recent evidence in human NPCs indicates that ATRX forms nuclear puncta with condensate\u2010like properties, associates with neurogenic enhancer\u2010rich regions, and incorporates regulatory factors such as CHD7 and p300. Perturbation of ATRX condensate formation is associated with changes in enhancer\u2010associated ATRX occupancy, neural gene\u2010expression programs, and neuroepithelial organization, suggesting a regulatory mode that may complement canonical heterochromatin\u2010associated functions. We propose a dual\u2010mode model in which folded domains contribute to chromatin anchoring at repressive regions, whereas intrinsically disordered regions support condensate\u2010associated organization at active developmental enhancers. We emphasize that whether ATRX condensates activate enhancers\n                    <jats:italic>de novo<\/jats:italic>\n                    , stabilize pre\u2010existing enhancer states, buffer transcriptional variability, or primarily organize cofactor localization remains unresolved. We also discuss limitations of the current evidence and outline acute, locus\u2010specific experiments needed to test the model.\n                  <\/jats:p>","DOI":"10.1002\/bies.70178","type":"journal-article","created":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T09:24:55Z","timestamp":1789118695000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["ATRX Condensates as Candidate Organizers of Enhancer\u2010Centered Nuclear Microenvironments in Neural Progenitors"],"prefix":"10.1002","volume":"48","author":[{"given":"Ryo","family":"Tomooka","sequence":"first","affiliation":[{"name":"Laboratory of Stem Cell Biology, Faculty of Human Sciences Waseda University  Tokorozawa Saitama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jun","family":"Kohyama","sequence":"additional","affiliation":[{"name":"Laboratory of Stem Cell Biology, Faculty of Human Sciences Waseda University  Tokorozawa Saitama Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,11]]},"reference":[{"key":"e_1_2_16_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg.2017.57"},{"key":"e_1_2_16_3_1","doi-asserted-by":"publisher","DOI":"10.3390\/genes6030641"},{"key":"e_1_2_16_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2016.02.007"},{"key":"e_1_2_16_5_1","doi-asserted-by":"publisher","DOI":"10.3389\/fncel.2014.00396"},{"key":"e_1_2_16_6_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.287.5457.1433"},{"key":"e_1_2_16_7_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aal2380"},{"key":"e_1_2_16_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/s44320\u2010024\u201000014\u2010z"},{"key":"e_1_2_16_9_1","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a020610"},{"key":"e_1_2_16_10_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3413"},{"key":"e_1_2_16_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3207"},{"key":"e_1_2_16_12_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3607"},{"key":"e_1_2_16_13_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00018\u2010016\u20102184\u20103"},{"key":"e_1_2_16_14_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2017.7"},{"key":"e_1_2_16_15_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aar3958"},{"key":"e_1_2_16_16_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aar4199"},{"key":"e_1_2_16_17_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.47098"},{"key":"e_1_2_16_18_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.331520.119"},{"key":"e_1_2_16_19_1","doi-asserted-by":"publisher","DOI":"10.3390\/genes14020499"},{"key":"e_1_2_16_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2010.09.023"},{"key":"e_1_2_16_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2010.01.003"},{"key":"e_1_2_16_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/0092\u20108674(95)90287\u20102"},{"key":"e_1_2_16_23_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature10833"},{"key":"e_1_2_16_24_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tig.2023.02.009"},{"key":"e_1_2_16_25_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010022\u201031194\u20107"},{"key":"e_1_2_16_26_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010025\u201061881\u20100"},{"key":"e_1_2_16_27_1","doi-asserted-by":"publisher","DOI":"10.1038\/ng.825"},{"key":"e_1_2_16_28_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.301887.117"},{"key":"e_1_2_16_29_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature07730"},{"key":"e_1_2_16_30_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1016071107"},{"key":"e_1_2_16_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2013.03.035"},{"key":"e_1_2_16_32_1","doi-asserted-by":"publisher","DOI":"10.1038\/nsmb.2062"},{"key":"e_1_2_16_33_1","doi-asserted-by":"publisher","DOI":"10.1038\/nsmb.2070"},{"key":"e_1_2_16_34_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tibs.2013.06.011"},{"key":"e_1_2_16_35_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010019\u20101464\u20100"},{"key":"e_1_2_16_36_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2017.02.007"},{"key":"e_1_2_16_37_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2025.11.026"},{"key":"e_1_2_16_38_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molcel.2019.07.009"},{"key":"e_1_2_16_39_1","doi-asserted-by":"publisher","DOI":"10.1101\/gr.086983.108"},{"key":"e_1_2_16_40_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau0320"},{"key":"e_1_2_16_41_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2016.76"},{"key":"e_1_2_16_42_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2013.10.037"},{"key":"e_1_2_16_43_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2727"},{"key":"e_1_2_16_44_1","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a008359"},{"key":"e_1_2_16_45_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2014.03.018"},{"key":"e_1_2_16_46_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2025.03.031"},{"key":"e_1_2_16_47_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkad045"},{"key":"e_1_2_16_48_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.TM118.001189"},{"key":"e_1_2_16_49_1","doi-asserted-by":"publisher","DOI":"10.15252\/embj.201696394"},{"key":"e_1_2_16_50_1","doi-asserted-by":"publisher","DOI":"10.3390\/ijms20215501"},{"key":"e_1_2_16_51_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2018.12.035"},{"key":"e_1_2_16_52_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41580\u2010020\u201000303\u2010z"},{"key":"e_1_2_16_53_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.03.001"},{"key":"e_1_2_16_54_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41589\u2010018\u20100021\u20108"},{"key":"e_1_2_16_55_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1417910112"},{"key":"e_1_2_16_56_1","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth763"},{"key":"e_1_2_16_57_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010024\u201050858\u20100"},{"key":"e_1_2_16_58_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.chembiol.2025.01.006"},{"key":"e_1_2_16_59_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molcel.2020.04.032"},{"key":"e_1_2_16_60_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010025\u201009141\u20105"},{"key":"e_1_2_16_61_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2016.11.054"},{"key":"e_1_2_16_62_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2018.10.048"},{"key":"e_1_2_16_63_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1399\u20100004.2006.00641.x"},{"key":"e_1_2_16_64_1","doi-asserted-by":"publisher","DOI":"10.1002\/humu.20734"},{"key":"e_1_2_16_65_1","doi-asserted-by":"crossref","unstructured":"J.Guez J. K.Goodrich M. A.Moldovan et\u00a0al. \u201cIntegrating 730 947 Exome Sequences With Clinical Literature Improves Gene Discovery \u201d preprint medRxiv March 25 2026 https:\/\/doi.org\/10.64898\/2026.03.23.26349081.","DOI":"10.64898\/2026.03.23.26349081"},{"key":"e_1_2_16_66_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.46883"},{"key":"e_1_2_16_67_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev\u2010genet\u2010112618\u2010043527"},{"key":"e_1_2_16_68_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010021\u201003662\u20105"},{"key":"e_1_2_16_69_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2018.08.003"},{"key":"e_1_2_16_70_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaw8653"},{"key":"e_1_2_16_71_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.R115.695056"},{"key":"e_1_2_16_72_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrm3920"},{"key":"e_1_2_16_73_1","doi-asserted-by":"publisher","DOI":"10.1021\/acs.biochem.7b01136"},{"key":"e_1_2_16_74_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41573\u2010025\u201001220\u20106"},{"key":"e_1_2_16_75_1","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.aac8228"},{"key":"e_1_2_16_76_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1207313"},{"key":"e_1_2_16_77_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010018\u201003476\u20106"},{"key":"e_1_2_16_78_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12967\u2010023\u201004082\u2010x"}],"container-title":["BioEssays"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/bies.70178","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/bies.70178","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/bies.70178","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T09:15:07Z","timestamp":1790673307000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/bies.70178"}},"subtitle":["A Hypothesis for Enhancer\u2010Associated ATRX Function in Neural Progenitors"],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":77,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2026,9]]}},"alternative-id":["10.1002\/bies.70178"],"URL":"https:\/\/doi.org\/10.1002\/bies.70178","archive":["Portico"],"relation":{},"ISSN":["0265-9247","1521-1878"],"issn-type":[{"value":"0265-9247","type":"print"},{"value":"1521-1878","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"2026-06-10","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-08-20","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-11","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70178"}}