{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T02:45:30Z","timestamp":1782873930994,"version":"3.54.5"},"reference-count":67,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T00:00:00Z","timestamp":1778112000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"vor","delay-in-days":55,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"Spanish Ministry of Science","award":["PID2023-146239OB-I00"],"award-info":[{"award-number":["PID2023-146239OB-I00"]}]},{"name":"Institut National du Cancer","award":["PRT-K16-071"],"award-info":[{"award-number":["PRT-K16-071"]}]},{"DOI":"10.13039\/100010665","name":"H2020 Marie Sk\u0142odowska-Curie Actions","doi-asserted-by":"publisher","award":["# 813091 ARCH"],"award-info":[{"award-number":["# 813091 ARCH"]}],"id":[{"id":"10.13039\/100010665","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Institute for Cancer and Immunology"},{"DOI":"10.13039\/501100014182","name":"Cancerop\u00f4le Provence Alpes C\u00f4te d\u2019Azur","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100014182","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Epigenetics &amp; Chromatin"],"DOI":"10.1186\/s13072-026-00679-w","type":"journal-article","created":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T18:44:49Z","timestamp":1778179489000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["H1.3 depletion in AML cells prompts H1.2 redistribution, chromatin remodeling and cell cycle defects"],"prefix":"10.1186","volume":"19","author":[{"given":"Clara","family":"Tellez-Quijorna","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lia","family":"N\u2019Guyen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"N\u00faria","family":"Serna-Pujol","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maeva","family":"Rodies","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pau","family":"Homs-Aguad\u00e9","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julien","family":"Vernerey","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Albert","family":"Jordan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Estelle","family":"Duprez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,7]]},"reference":[{"key":"679_CR1","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1038\/s41392-024-02030-9","volume":"9","author":"Y Chen","year":"2024","unstructured":"Chen Y, Liang R, Li Y, Jiang L, Ma D, Luo Q, et al. Chromatin accessibility: biological functions, molecular mechanisms and therapeutic application. Sig Transduct Target Ther. 2024;9:340. https:\/\/doi.org\/10.1038\/s41392-024-02030-9.","journal-title":"Sig Transduct Target Ther"},{"key":"679_CR2","doi-asserted-by":"publisher","first-page":"737","DOI":"10.1038\/s41576-020-0270-8","volume":"21","author":"Z Dai","year":"2020","unstructured":"Dai Z, Ramesh V, Locasale JW. The evolving metabolic landscape of chromatin biology and epigenetics. Nat Rev Genet. 2020;21:737\u201353. https:\/\/doi.org\/10.1038\/s41576-020-0270-8.","journal-title":"Nat Rev Genet"},{"key":"679_CR3","doi-asserted-by":"publisher","first-page":"868","DOI":"10.1126\/science.184.4139.868","volume":"184","author":"RD Kornberg","year":"1974","unstructured":"Kornberg RD, Chromatin Structure. A Repeating Unit of Histones and DNA: Chromatin structure is based on a repeating unit of eight histone molecules and about 200 DNA base pairs. Science. 1974;184:868\u201371. https:\/\/doi.org\/10.1126\/science.184.4139.868.","journal-title":"Science"},{"key":"679_CR4","doi-asserted-by":"publisher","first-page":"1298","DOI":"10.1038\/ncb3629","volume":"19","author":"MA Reid","year":"2017","unstructured":"Reid MA, Dai Z, Locasale JW. The impact of cellular metabolism on chromatin dynamics and epigenetics. Nat Cell Biol. 2017;19:1298\u2013306. https:\/\/doi.org\/10.1038\/ncb3629.","journal-title":"Nat Cell Biol"},{"key":"679_CR5","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1006\/jsbi.2000.4310","volume":"132","author":"G Wanner","year":"2000","unstructured":"Wanner G, Formanek H. A New Chromosome Model. J Struct Biol. 2000;132:147\u201361. https:\/\/doi.org\/10.1006\/jsbi.2000.4310.","journal-title":"J Struct Biol"},{"key":"679_CR6","doi-asserted-by":"publisher","first-page":"11170","DOI":"10.1073\/pnas.92.24.11170","volume":"92","author":"G Arents","year":"1995","unstructured":"Arents G, Moudrianakis EN. The histone fold: a ubiquitous architectural motif utilized in DNA compaction and protein dimerization. Proc Natl Acad Sci USA. 1995;92:11170\u20134. https:\/\/doi.org\/10.1073\/pnas.92.24.11170.","journal-title":"Proc Natl Acad Sci USA"},{"key":"679_CR7","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/0092-8674(75)90149-X","volume":"4","author":"P Oudet","year":"1975","unstructured":"Oudet P, Gross-Bellard M, Chambon P. Electron microscopic and biochemical evidence that chromatin structure is a repeating unit. Cell. 1975;4:281\u2013300. https:\/\/doi.org\/10.1016\/0092-8674(75)90149-X.","journal-title":"Cell"},{"key":"679_CR8","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1016\/S0022-2836(77)80019-3","volume":"109","author":"M Noll","year":"1977","unstructured":"Noll M, Kornberg RD. Action of micrococcal nuclease on chromatin and the location of histone H1. J Mol Biol. 1977;109:393\u2013404. https:\/\/doi.org\/10.1016\/S0022-2836(77)80019-3.","journal-title":"J Mol Biol"},{"key":"679_CR9","doi-asserted-by":"publisher","first-page":"691","DOI":"10.1016\/0092-8674(78)90219-2","volume":"13","author":"RT Simpson","year":"1978","unstructured":"Simpson RT. Structure of chromatin containing extensively acetylated H3 and H4. Cell. 1978;13:691\u20139. https:\/\/doi.org\/10.1016\/0092-8674(78)90219-2.","journal-title":"Cell"},{"key":"679_CR10","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1038\/nrm.2017.94","volume":"19","author":"DV Fyodorov","year":"2018","unstructured":"Fyodorov DV, Zhou B-R, Skoultchi AI, Bai Y. Emerging roles of linker histones in regulating chromatin structure and function. Nat Rev Mol Cell Biol. 2018;19:192\u2013206. https:\/\/doi.org\/10.1038\/nrm.2017.94.","journal-title":"Nat Rev Mol Cell Biol"},{"key":"679_CR11","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.gene.2008.11.003","volume":"431","author":"N Happel","year":"2009","unstructured":"Happel N, Doenecke D. Histone H1 and its isoforms: Contribution to chromatin structure and function. Gene. 2009;431:1\u201312. https:\/\/doi.org\/10.1016\/j.gene.2008.11.003.","journal-title":"Gene"},{"key":"679_CR12","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1101\/gad.344531.120","volume":"35","author":"L Prendergast","year":"2021","unstructured":"Prendergast L, Reinberg D. The missing link er: emerging trends for H1 variant-specific functions. Genes Dev. 2021;35:40\u201358. https:\/\/doi.org\/10.1101\/gad.344531.120.","journal-title":"Genes Dev"},{"key":"679_CR13","doi-asserted-by":"publisher","first-page":"2142","DOI":"10.1016\/j.celrep.2013.05.003","volume":"3","author":"A Izzo","year":"2013","unstructured":"Izzo A, Kamieniarz-Gdula K, Ram\u00edrez F, Noureen N, Kind J, Manke T, et al. The Genomic Landscape of the Somatic Linker Histone Subtypes H1.1 to H1.5 in Human Cells. Cell Rep. 2013;3:2142\u201354. https:\/\/doi.org\/10.1016\/j.celrep.2013.05.003.","journal-title":"Cell Rep"},{"key":"679_CR14","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/1756-8935-5-7","volume":"5","author":"PB Talbert","year":"2012","unstructured":"Talbert PB, Ahmad K, Almouzni G, Ausi\u00f3 J, Berger F, Bhalla PL, et al. A unified phylogeny-based nomenclature for histone variants. Epigenetics Chromatin. 2012;5:7. https:\/\/doi.org\/10.1186\/1756-8935-5-7.","journal-title":"Epigenetics Chromatin"},{"key":"679_CR15","doi-asserted-by":"publisher","first-page":"e1000227","DOI":"10.1371\/journal.pgen.1000227","volume":"4","author":"M Sancho","year":"2008","unstructured":"Sancho M, Diani E, Beato M, Jordan A. Depletion of Human Histone H1 Variants Uncovers Specific Roles in Gene Expression and Cell Growth. PLoS Genet. 2008;4:e1000227. . https:\/\/doi.org\/10.1371\/journal.pgen.1000227","journal-title":"PLoS Genet"},{"key":"679_CR16","doi-asserted-by":"publisher","first-page":"773195","DOI":"10.3389\/fcell.2021.773195","volume":"9","author":"S Lai","year":"2022","unstructured":"Lai S, Jia J, Cao X, Zhou P-K, Gao S. Molecular and Cellular Functions of the Linker Histone H1.2. Front Cell Dev Biol. 2022;9:773195. https:\/\/doi.org\/10.3389\/fcell.2021.773195.","journal-title":"Front Cell Dev Biol"},{"key":"679_CR17","doi-asserted-by":"publisher","first-page":"1793","DOI":"10.1093\/nar\/gkae014","volume":"52","author":"M Salinas-Pena","year":"2024","unstructured":"Salinas-Pena M, Serna-Pujol N, Jordan A. Genomic profiling of six human somatic histone H1 variants denotes that H1X accumulates at recently incorporated transposable elements. Nucleic Acids Res. 2024;52:1793\u2013813. https:\/\/doi.org\/10.1093\/nar\/gkae014.","journal-title":"Nucleic Acids Res"},{"key":"679_CR18","doi-asserted-by":"publisher","first-page":"1008","DOI":"10.7150\/ijbs.19891","volume":"13","author":"X Ye","year":"2017","unstructured":"Ye X, Feng C, Gao T, Mu G, Zhu W, Yang Y. Linker Histone in Diseases. Int J Biol Sci. 2017;13:1008\u201318. https:\/\/doi.org\/10.7150\/ijbs.19891.","journal-title":"Int J Biol Sci"},{"key":"679_CR19","doi-asserted-by":"publisher","first-page":"810","DOI":"10.1186\/s12885-017-3834-z","volume":"17","author":"M Bauden","year":"2017","unstructured":"Bauden M, Kristl T, Sasor A, Andersson B, Marko-Varga G, Andersson R, et al. Histone profiling reveals the H1.3 histone variant as a prognostic biomarker for pancreatic ductal adenocarcinoma. BMC Cancer. 2017;17:810. https:\/\/doi.org\/10.1186\/s12885-017-3834-z.","journal-title":"BMC Cancer"},{"key":"679_CR20","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1038\/s41525-023-00367-8","volume":"8","author":"ER Bonner","year":"2023","unstructured":"Bonner ER, Dawood A, Gordish-Dressman H, Eze A, Bhattacharya S, Yadavilli S, et al. Pan-cancer atlas of somatic core and linker histone mutations. npj Genom Med. 2023;8:23. https:\/\/doi.org\/10.1038\/s41525-023-00367-8.","journal-title":"npj Genom Med"},{"key":"679_CR21","doi-asserted-by":"publisher","first-page":"1202","DOI":"10.1038\/leu.2014.339","volume":"29","author":"G Tiberi","year":"2015","unstructured":"Tiberi G, Pekowska A, Oudin C, Ivey A, Autret A, Prebet T, et al. PcG methylation of the HIST1 cluster defines an epigenetic marker of acute myeloid leukemia. Leukemia. 2015;29:1202\u20136. https:\/\/doi.org\/10.1038\/leu.2014.339.","journal-title":"Leukemia"},{"key":"679_CR22","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1186\/s13148-019-0738-6","volume":"11","author":"S Garciaz","year":"2019","unstructured":"Garciaz S, N\u2019guyen Dasi L, Finetti P, Chevalier C, Vernerey J, Poplineau M, et al. Epigenetic down-regulation of the HIST1 locus predicts better prognosis in acute myeloid leukemia with NPM1 mutation. Clin Epigenet. 2019;11:141. https:\/\/doi.org\/10.1186\/s13148-019-0738-6.","journal-title":"Clin Epigenet"},{"key":"679_CR23","doi-asserted-by":"publisher","first-page":"3047","DOI":"10.1093\/bioinformatics\/btw354","volume":"32","author":"P Ewels","year":"2016","unstructured":"Ewels P, Magnusson M, Lundin S, K\u00e4ller M. MultiQC: summarize analysis results for multiple tools and samples in a single report. Bioinformatics. 2016;32:3047\u20138. https:\/\/doi.org\/10.1093\/bioinformatics\/btw354.","journal-title":"Bioinformatics"},{"key":"679_CR24","doi-asserted-by":"publisher","first-page":"e108","DOI":"10.1093\/nar\/gkt214","volume":"41","author":"Y Liao","year":"2013","unstructured":"Liao Y, Smyth GK, Shi W. The Subread aligner: fast, accurate and scalable read mapping by seed-and-vote. Nucleic Acids Res. 2013;41:e108\u2013108. https:\/\/doi.org\/10.1093\/nar\/gkt214.","journal-title":"Nucleic Acids Res"},{"key":"679_CR25","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1093\/bioinformatics\/btt656","volume":"30","author":"Y Liao","year":"2014","unstructured":"Liao Y, Smyth GK, Shi W. featureCounts: an efficient general purpose program for assigning sequence reads to genomic features. Bioinformatics. 2014;30:923\u201330. https:\/\/doi.org\/10.1093\/bioinformatics\/btt656.","journal-title":"Bioinformatics"},{"key":"679_CR26","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","volume":"15","author":"MI Love","year":"2014","unstructured":"Love MI, Huber W, Anders S. Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2. Genome Biol. 2014;15:550. https:\/\/doi.org\/10.1186\/s13059-014-0550-8.","journal-title":"Genome Biol"},{"key":"679_CR27","doi-asserted-by":"publisher","first-page":"W160","DOI":"10.1093\/nar\/gkw257","volume":"44","author":"F Ram\u00edrez","year":"2016","unstructured":"Ram\u00edrez F, Ryan DP, Gr\u00fcning B, Bhardwaj V, Kilpert F, Richter AS, et al. deepTools2: a next generation web server for deep-sequencing data analysis. Nucleic Acids Res. 2016;44:W160\u20135. https:\/\/doi.org\/10.1093\/nar\/gkw257.","journal-title":"Nucleic Acids Res"},{"key":"679_CR28","doi-asserted-by":"publisher","first-page":"11622","DOI":"10.1093\/nar\/gkx746","volume":"45","author":"A Izquierdo-Bouldstridge","year":"2017","unstructured":"Izquierdo-Bouldstridge A, Bustillos A, Bonet-Costa C, Aribau-Miralb\u00e9s P, Garc\u00eda-Gomis D, Dabad M, et al. Histone H1 depletion triggers an interferon response in cancer cells via activation of heterochromatic repeats. Nucleic Acids Res. 2017;45:11622\u201342. https:\/\/doi.org\/10.1093\/nar\/gkx746.","journal-title":"Nucleic Acids Res"},{"key":"679_CR29","doi-asserted-by":"publisher","first-page":"W501","DOI":"10.1093\/nar\/gkx372","volume":"45","author":"W Luo","year":"2017","unstructured":"Luo W, Pant G, Bhavnasi YK, Blanchard SG, Brouwer C. Pathview Web: user friendly pathway visualization and data integration. Nucleic Acids Res. 2017;45:W501\u20138. https:\/\/doi.org\/10.1093\/nar\/gkx372.","journal-title":"Nucleic Acids Res"},{"key":"679_CR30","doi-asserted-by":"publisher","first-page":"3339","DOI":"10.1093\/nar\/gky080","volume":"46","author":"M Koubi","year":"2018","unstructured":"Koubi M, Poplineau M, Vernerey J, N\u2019Guyen L, Tiberi G, Garciaz S, et al. Regulation of the positive transcriptional effect of PLZF through a non-canonical EZH2 activity. Nucleic Acids Res. 2018;46:3339\u201350. https:\/\/doi.org\/10.1093\/nar\/gky080.","journal-title":"Nucleic Acids Res"},{"key":"679_CR31","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1038\/nmeth.1923","volume":"9","author":"B Langmead","year":"2012","unstructured":"Langmead B, Salzberg SL. Fast gapped-read alignment with Bowtie 2. Nat Methods. 2012;9:357\u20139. https:\/\/doi.org\/10.1038\/nmeth.1923.","journal-title":"Nat Methods"},{"key":"679_CR32","doi-asserted-by":"publisher","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","volume":"25","author":"H Li","year":"2009","unstructured":"Li H, Handsaker B, Wysoker A, Fennell T, Ruan J, Homer N, et al. The Sequence Alignment\/Map format and SAMtools. Bioinformatics. 2009;25:2078\u20139. https:\/\/doi.org\/10.1093\/bioinformatics\/btp352.","journal-title":"Bioinformatics"},{"key":"679_CR33","doi-asserted-by":"publisher","first-page":"841","DOI":"10.1093\/bioinformatics\/btq033","volume":"26","author":"AR Quinlan","year":"2010","unstructured":"Quinlan AR, Hall IM. BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics. 2010;26:841\u20132. https:\/\/doi.org\/10.1093\/bioinformatics\/btq033.","journal-title":"Bioinformatics"},{"key":"679_CR34","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1038\/nbt.1754","volume":"29","author":"JT Robinson","year":"2011","unstructured":"Robinson JT, Thorvaldsd\u00f3ttir H, Winckler W, Guttman M, Lander ES, Getz G, et al. Integrative genomics viewer. Nat Biotechnol. 2011;29:24\u20136. https:\/\/doi.org\/10.1038\/nbt.1754.","journal-title":"Nat Biotechnol"},{"key":"679_CR35","doi-asserted-by":"publisher","first-page":"1989","DOI":"10.1111\/febs.15549","volume":"288","author":"N Serna-Pujol","year":"2021","unstructured":"Serna-Pujol N, Salinas\u2010Pena M, Mugianesi F, Lopez\u2010Anguita N, Torrent\u2010Llagostera F, Izquierdo\u2010Bouldstridge A, et al. TADs enriched in histone H1.2 strongly overlap with the B compartment, inaccessible chromatin, and AT\u2010rich Giemsa bands. FEBS J. 2021;288:1989\u20132013. https:\/\/doi.org\/10.1111\/febs.15549.","journal-title":"FEBS J"},{"key":"679_CR36","doi-asserted-by":"publisher","first-page":"R137","DOI":"10.1186\/gb-2008-9-9-r137","volume":"9","author":"Y Zhang","year":"2008","unstructured":"Zhang Y, Liu T, Meyer CA, Eeckhoute J, Johnson DS, Bernstein BE, et al. Model-based Analysis of ChIP-Seq (MACS). Genome Biol. 2008;9:R137. https:\/\/doi.org\/10.1186\/gb-2008-9-9-r137.","journal-title":"Genome Biol"},{"key":"679_CR37","doi-asserted-by":"publisher","first-page":"3593","DOI":"10.1093\/bioinformatics\/btv422","volume":"31","author":"Y Jin","year":"2015","unstructured":"Jin Y, Tam OH, Paniagua E, Hammell M. TEtranscripts: a package for including transposable elements in differential expression analysis of RNA-seq datasets. Bioinformatics. 2015;31:3593\u20139. https:\/\/doi.org\/10.1093\/bioinformatics\/btv422.","journal-title":"Bioinformatics"},{"key":"679_CR38","doi-asserted-by":"publisher","first-page":"9354","DOI":"10.1038\/s41598-019-45839-z","volume":"9","author":"HM Amemiya","year":"2019","unstructured":"Amemiya HM, Kundaje A, Boyle AP. The ENCODE Blacklist: Identification of Problematic Regions of the Genome. Sci Rep. 2019;9:9354. https:\/\/doi.org\/10.1038\/s41598-019-45839-z.","journal-title":"Sci Rep"},{"key":"679_CR39","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1016\/j.molcel.2010.05.004","volume":"38","author":"S Heinz","year":"2010","unstructured":"Heinz S, Benner C, Spann N, Bertolino E, Lin YC, Laslo P, et al. Simple Combinations of Lineage-Determining Transcription Factors Prime cis-Regulatory Elements Required for Macrophage and B Cell Identities. Mol Cell. 2010;38:576\u201389. https:\/\/doi.org\/10.1016\/j.molcel.2010.05.004.","journal-title":"Mol Cell"},{"key":"679_CR40","doi-asserted-by":"publisher","first-page":"e585","DOI":"10.1002\/cpz1.585","volume":"2","author":"Q Wang","year":"2022","unstructured":"Wang Q, Li M, Wu T, Zhan L, Li L, Chen M, et al. Exploring Epigenomic Datasets by ChIPseeker. Curr Protocols. 2022;2:e585. https:\/\/doi.org\/10.1002\/cpz1.585.","journal-title":"Curr Protocols"},{"key":"679_CR41","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1007\/s004180000146","volume":"113","author":"B Drabent","year":"2000","unstructured":"Drabent B, Saftig P, Bode C, Doenecke D. Spermatogenesis proceeds normally in mice without linker histone H1t. Histochem Cell Biol. 2000;113:433\u201342. https:\/\/doi.org\/10.1007\/s004180000146.","journal-title":"Histochem Cell Biol"},{"key":"679_CR42","doi-asserted-by":"publisher","first-page":"7933","DOI":"10.1128\/MCB.21.23.7933-7943.2001","volume":"21","author":"Y Fan","year":"2001","unstructured":"Fan Y, Sirotkin A, Russell RG, Ayala J, Skoultchi AI. Individual Somatic H1 Subtypes Are Dispensable for Mouse Development Even in Mice Lacking the H10 Replacement Subtype. Mol Cell Biol. 2001;21:7933\u201343. https:\/\/doi.org\/10.1128\/MCB.21.23.7933-7943.2001.","journal-title":"Mol Cell Biol"},{"key":"679_CR43","doi-asserted-by":"publisher","first-page":"14251","DOI":"10.1073\/pnas.1920725117","volume":"117","author":"SE Healton","year":"2020","unstructured":"Healton SE, Pinto HD, Mishra LN, Hamilton GA, Wheat JC, Swist-Rosowska K, et al. H1 linker histones silence repetitive elements by promoting both histone H3K9 methylation and chromatin compaction. Proc Natl Acad Sci USA. 2020;117:14251\u20138. https:\/\/doi.org\/10.1073\/pnas.1920725117.","journal-title":"Proc Natl Acad Sci USA"},{"key":"679_CR44","doi-asserted-by":"publisher","first-page":"4081","DOI":"10.1038\/s41467-023-39846-y","volume":"14","author":"C Liu","year":"2023","unstructured":"Liu C, Yu J, Song A, Wang M, Hu J, Chen P, et al. Histone H1 facilitates restoration of H3K27me3 during DNA replication by chromatin compaction. Nat Commun. 2023;14:4081. https:\/\/doi.org\/10.1038\/s41467-023-39846-y.","journal-title":"Nat Commun"},{"key":"679_CR45","doi-asserted-by":"publisher","first-page":"819","DOI":"10.1126\/science.1149250","volume":"319","author":"R Krishnakumar","year":"2008","unstructured":"Krishnakumar R, Gamble MJ, Frizzell KM, Berrocal JG, Kininis M, Kraus WL. Reciprocal Binding of PARP-1 and Histone H1 at Promoters Specifies Transcriptional Outcomes. Science. 2008;319:819\u201321. https:\/\/doi.org\/10.1126\/science.1149250.","journal-title":"Science"},{"key":"679_CR46","doi-asserted-by":"publisher","first-page":"3801","DOI":"10.1016\/j.molcel.2023.10.003","volume":"83","author":"C Pascal","year":"2023","unstructured":"Pascal C, Zonszain J, Hameiri O, Gargi-Levi C, Lev-Maor G, Tammer L, et al. Human histone H1 variants impact splicing outcome by controlling RNA polymerase II elongation. Mol Cell. 2023;83:3801\u2013e38178. https:\/\/doi.org\/10.1016\/j.molcel.2023.10.003.","journal-title":"Mol Cell"},{"key":"679_CR47","doi-asserted-by":"publisher","first-page":"16714","DOI":"10.1038\/srep16714","volume":"5","author":"J-M Kim","year":"2015","unstructured":"Kim J-M, Kim K, Punj V, Liang G, Ulmer TS, Lu W, et al. Linker histone H1.2 establishes chromatin compaction and gene silencing through recognition of H3K27me3. Sci Rep. 2015;5:16714. https:\/\/doi.org\/10.1038\/srep16714.","journal-title":"Sci Rep"},{"key":"679_CR48","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1016\/j.molcel.2019.10.011","volume":"77","author":"J Choi","year":"2020","unstructured":"Choi J, Lyons DB, Kim MY, Moore JD, Zilberman D. DNA Methylation and Histone H1 Jointly Repress Transposable Elements and Aberrant Intragenic Transcripts. Mol Cell. 2020;77:310\u2013e3237. https:\/\/doi.org\/10.1016\/j.molcel.2019.10.011.","journal-title":"Mol Cell"},{"key":"679_CR49","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1007\/978-1-0716-2736-5_5","volume":"2579","author":"RC Wang","year":"2022","unstructured":"Wang RC, Wang Z. Synchronization of Cultured Cells to G1, S, G2, and M Phases by Double Thymidine Block. Methods Mol Biol. 2022;2579:61\u201371. https:\/\/doi.org\/10.1007\/978-1-0716-2736-5_5.","journal-title":"Methods Mol Biol"},{"key":"679_CR50","doi-asserted-by":"publisher","first-page":"e1003417","DOI":"10.1371\/journal.pgen.1003417","volume":"9","author":"K Cao","year":"2013","unstructured":"Cao K, Lailler N, Zhang Y, Kumar A, Uppal K, Liu Z, et al. High-Resolution Mapping of H1 Linker Histone Variants in Embryonic Stem Cells. PLoS Genet. 2013;9:e1003417. . https:\/\/doi.org\/10.1371\/journal.pgen.1003417","journal-title":"PLoS Genet"},{"key":"679_CR51","doi-asserted-by":"publisher","first-page":"4474","DOI":"10.1093\/nar\/gku079","volume":"42","author":"L Mill\u00e1n-Ari\u00f1o","year":"2014","unstructured":"Mill\u00e1n-Ari\u00f1o L, Islam ABMMK, Izquierdo-Bouldstridge A, Mayor R, Terme J-M, Luque N, et al. Mapping of six somatic linker histone H1 variants in human breast cancer cells uncovers specific features of H1.2. Nucleic Acids Res. 2014;42:4474\u201393. https:\/\/doi.org\/10.1093\/nar\/gku079.","journal-title":"Nucleic Acids Res"},{"key":"679_CR52","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1038\/s41586-020-3032-z","volume":"589","author":"MA Willcockson","year":"2021","unstructured":"Willcockson MA, Healton SE, Weiss CN, Bartholdy BA, Botbol Y, Mishra LN, et al. H1 histones control the epigenetic landscape by local chromatin compaction. Nature. 2021;589:293\u20138. https:\/\/doi.org\/10.1038\/s41586-020-3032-z.","journal-title":"Nature"},{"key":"679_CR53","doi-asserted-by":"publisher","first-page":"2503","DOI":"10.1016\/j.molcel.2025.05.031","volume":"85","author":"RE Matthews","year":"2025","unstructured":"Matthews RE, Danac JMC, Naden EL, Farleigh Smith LE, Lestari S, Gungi A, et al. CRAMP1 drives linker histone expression to enable Polycomb repression. Mol Cell. 2025;85:2503\u2013e25168. https:\/\/doi.org\/10.1016\/j.molcel.2025.05.031.","journal-title":"Mol Cell"},{"key":"679_CR54","doi-asserted-by":"publisher","first-page":"826","DOI":"10.1016\/j.pathol.2024.03.012","volume":"56","author":"AG Lashen","year":"2024","unstructured":"Lashen AG, Almalki N, Toss M, Mirza S, Malki MI, Rutland CS, et al. The characteristics and prognostic significance of histone H1 expression in breast cancer. Pathology. 2024;56:826\u201333. https:\/\/doi.org\/10.1016\/j.pathol.2024.03.012.","journal-title":"Pathology"},{"key":"679_CR55","doi-asserted-by":"publisher","first-page":"3892","DOI":"10.1093\/nar\/gkac226","volume":"50","author":"N Serna-Pujol","year":"2022","unstructured":"Serna-Pujol N, Salinas-Pena M, Mugianesi F, Le Dily F, Marti-Renom MA, Jordan A. Coordinated changes in gene expression, H1 variant distribution and genome 3D conformation in response to H1 depletion. Nucleic Acids Res. 2022;50:3892\u2013910. https:\/\/doi.org\/10.1093\/nar\/gkac226.","journal-title":"Nucleic Acids Res"},{"key":"679_CR56","doi-asserted-by":"publisher","first-page":"114137","DOI":"10.1016\/j.celrep.2024.114137","volume":"43","author":"P Bujosa","year":"2024","unstructured":"Bujosa P, Reina O, Caball\u00e9 A, Casas-Lamesa A, Torras-Llort M, P\u00e9rez-Rold\u00e1n J, et al. Linker histone H1 regulates homeostasis of heterochromatin-associated cRNAs. Cell Rep. 2024;43:114137. https:\/\/doi.org\/10.1016\/j.celrep.2024.114137.","journal-title":"Cell Rep"},{"key":"679_CR57","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1038\/emboj.2011.399","volume":"31","author":"MD Lynch","year":"2012","unstructured":"Lynch MD, Smith AJH, De Gobbi M, Flenley M, Hughes JR, Vernimmen D, et al. An interspecies analysis reveals a key role for unmethylated CpG dinucleotides in vertebrate Polycomb complex recruitment. EMBO J. 2012;31:317\u201329. https:\/\/doi.org\/10.1038\/emboj.2011.399.","journal-title":"EMBO J"},{"key":"679_CR58","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1001244","author":"EM Mendenhall","year":"2010","unstructured":"Mendenhall EM, Koche RP, Truong T, Zhou VW, Issac B, Chi AS, et al. GC-Rich Sequence Elements Recruit PRC2 in Mammalian ES Cells. PLoS Genet. 2010. . https:\/\/doi.org\/10.1371\/journal.pgen.1001244","journal-title":"PLoS Genet"},{"key":"679_CR59","doi-asserted-by":"publisher","first-page":"RP91306","DOI":"10.7554\/eLife.91306","volume":"12","author":"M Salinas-Pena","year":"2024","unstructured":"Salinas-Pena M, Rebollo E, Jordan A. Imaging analysis of six human histone H1 variants reveals universal enrichment of H1.2, H1.3, and H1.5 at the nuclear periphery and nucleolar H1X presence. Elife. 2024;12:RP91306. https:\/\/doi.org\/10.7554\/eLife.91306.","journal-title":"Elife"},{"key":"679_CR60","doi-asserted-by":"publisher","first-page":"375","DOI":"10.3390\/ijms26010375","volume":"26","author":"O Vlasova","year":"2025","unstructured":"Vlasova O, Antonova I, Magomedova K, Osipova A, Shtompel P, Borunova A, et al. Anticancer Plant Secondary Metabolites Evicting Linker Histone H1.2 from Chromatin Activate Type I Interferon Signaling. IJMS. 2025;26:375. https:\/\/doi.org\/10.3390\/ijms26010375.","journal-title":"IJMS"},{"key":"679_CR61","doi-asserted-by":"publisher","first-page":"6463","DOI":"10.1158\/0008-5472.CAN-13-2922","volume":"74","author":"M Medrzycki","year":"2014","unstructured":"Medrzycki M, Zhang Y, Zhang W, Cao K, Pan C, Lailler N, et al. Histone H1.3 Suppresses H19 Noncoding RNA Expression and Cell Growth of Ovarian Cancer Cells. Cancer Res. 2014;74:6463\u201373. https:\/\/doi.org\/10.1158\/0008-5472.CAN-13-2922.","journal-title":"Cancer Res"},{"key":"679_CR62","doi-asserted-by":"publisher","first-page":"326","DOI":"10.1038\/s41568-019-0143-7","volume":"19","author":"LN Kent","year":"2019","unstructured":"Kent LN, Leone G. The broken cycle: E2F dysfunction in cancer. Nat Rev Cancer. 2019;19:326\u201338. https:\/\/doi.org\/10.1038\/s41568-019-0143-7.","journal-title":"Nat Rev Cancer"},{"key":"679_CR63","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1038\/s41594-023-00942-8","volume":"30","author":"A Bujarrabal-Dueso","year":"2023","unstructured":"Bujarrabal-Dueso A, Sendtner G, Meyer DH, Chatzinikolaou G, Stratigi K, Garinis GA, et al. The DREAM complex functions as conserved master regulator of somatic DNA-repair capacities. Nat Struct Mol Biol. 2023;30:475\u201388. https:\/\/doi.org\/10.1038\/s41594-023-00942-8.","journal-title":"Nat Struct Mol Biol"},{"key":"679_CR64","doi-asserted-by":"publisher","first-page":"2193","DOI":"10.1016\/j.celrep.2017.05.053","volume":"19","author":"S Munro","year":"2017","unstructured":"Munro S, Hookway ES, Floderer M, Carr SM, Konietzny R, Kessler BM, et al. Linker Histone H1.2 Directs Genome-wide Chromatin Association of the Retinoblastoma Tumor Suppressor Protein and Facilitates Its Function. Cell Rep. 2017;19:2193\u2013201. https:\/\/doi.org\/10.1016\/j.celrep.2017.05.053.","journal-title":"Cell Rep"},{"key":"679_CR65","doi-asserted-by":"publisher","first-page":"1590","DOI":"10.1038\/s41467-018-03539-8","volume":"9","author":"R Almeida","year":"2018","unstructured":"Almeida R, Fern\u00e1ndez-Justel JM, Santa-Mar\u00eda C, Cadoret J-C, Cano-Aroca L, Lombra\u00f1a R, et al. Chromatin conformation regulates the coordination between DNA replication and transcription. Nat Commun. 2018;9:1590. https:\/\/doi.org\/10.1038\/s41467-018-03539-8.","journal-title":"Nat Commun"},{"key":"679_CR66","doi-asserted-by":"publisher","first-page":"e202301933","DOI":"10.26508\/lsa.202301933","volume":"6","author":"M Ozgencil","year":"2023","unstructured":"Ozgencil M, Dullovi A, Christiane Higos RC, Ho\u0159ej\u0161\u00ed Z, Bellelli R. The linker histone H1\u2013BRCA1 axis is a crucial mediator of replication fork stability. Life Sci Alliance. 2023;6:e202301933. https:\/\/doi.org\/10.26508\/lsa.202301933.","journal-title":"Life Sci Alliance"},{"key":"679_CR67","doi-asserted-by":"publisher","first-page":"2487","DOI":"10.1016\/j.molcel.2025.04.006","volume":"85","author":"A Ingham","year":"2025","unstructured":"Ingham A, de Vega IA, Morlot L, Gittens W, Hendriks IA, Kakulidis ES, et al. CRAMP1-dependent histone H1 biogenesis is essential for topoisomerase II inhibitor tolerance. Mol Cell. 2025;85:2487\u2013e250212. https:\/\/doi.org\/10.1016\/j.molcel.2025.04.006.","journal-title":"Mol Cell"}],"container-title":["Epigenetics &amp; Chromatin"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13072-026-00679-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13072-026-00679-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13072-026-00679-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T02:08:32Z","timestamp":1782871712000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s13072-026-00679-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,7]]},"references-count":67,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["679"],"URL":"https:\/\/doi.org\/10.1186\/s13072-026-00679-w","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.07.28.667197","asserted-by":"object"}]},"ISSN":["1756-8935"],"issn-type":[{"value":"1756-8935","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,7]]},"assertion":[{"value":"9 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 April 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"28"}}