{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:35:15Z","timestamp":1791246915306,"version":"4.1.0"},"reference-count":111,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2019,10,22]],"date-time":"2019-10-22T00:00:00Z","timestamp":1571702400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["PJT-162120"],"award-info":[{"award-number":["PJT-162120"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"DOI":"10.3389\/fcell.2019.00241","type":"journal-article","created":{"date-parts":[[2019,10,22]],"date-time":"2019-10-22T04:22:35Z","timestamp":1571718155000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":102,"title":["Escape From X-Chromosome Inactivation: An Evolutionary Perspective"],"prefix":"10.3389","volume":"7","author":[{"given":"Bronwyn J.","family":"Posynick","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carolyn J.","family":"Brown","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2019,10,22]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"963","DOI":"10.1038\/415963a","article-title":"Human spermatozoa: the future of sex.","volume":"415","author":"Aitken","year":"2002","journal-title":"Nature"},{"key":"B2","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1007\/s00412-011-0343-8","article-title":"A cross-species comparison of escape from X inactivation in Eutheria: implications for evolution of X chromosome inactivation.","volume":"121","author":"Al Nadaf","year":"2011","journal-title":"Chromosoma"},{"key":"B3","doi-asserted-by":"publisher","first-page":"350","DOI":"10.1016\/j.tig.2011.05.005","article-title":"Are all sex chromosomes created equal?","volume":"27","author":"Bachtrog","year":"2011","journal-title":"Trends Genet."},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001899","article-title":"Sex determination: why so many ways of doing it?","volume":"12","author":"Bachtrog","year":"2014","journal-title":"PLoS Biol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"6634","DOI":"10.1073\/pnas.97.12.6634","article-title":"Molecular evidence for a relationship between LINE-1 elements and X chromosome inactivation: the Lyon repeat hypothesis.","volume":"97","author":"Bailey","year":"2000","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B6","doi-asserted-by":"publisher","first-page":"1224","DOI":"10.1093\/hmg\/ddt553","article-title":"DNA methylation profiling in X;autosome translocations supports a role for L1 repeats in the spread of X chromosome inactivation.","volume":"23","author":"Bala Tannan","year":"2014","journal-title":"Hum. Mol. Genet."},{"key":"B7","doi-asserted-by":"publisher","first-page":"348","DOI":"10.1016\/j.tig.2016.03.007","article-title":"Escape artists of the X chromosome.","volume":"32","author":"Balaton","year":"2016","journal-title":"Trends Genet."},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1186\/s13293-015-0053-7","article-title":"Derivation of consensus inactivation status for X-linked genes from genome-wide studies.","volume":"6","author":"Balaton","year":"2015","journal-title":"Biol. Sex Differ."},{"key":"B9","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1038\/nature13206","article-title":"Mammalian Y chromosomes retain widely expressed dosage-sensitive regulators.","volume":"508","author":"Bellott","year":"2014","journal-title":"Nature"},{"key":"B10","doi-asserted-by":"publisher","first-page":"612","DOI":"10.1038\/nature09172","article-title":"Convergent evolution of chicken Z and human X chromosomes by expansion and gene acquisition.","volume":"466","author":"Bellott","year":"2010","journal-title":"Nature"},{"key":"B11","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1016\/j.tree.2008.09.010","article-title":"The evolution of restricted recombination in sex chromosomes.","volume":"24","author":"Bergero","year":"2009","journal-title":"Trends Ecol. Evol."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1005079","article-title":"Escape from X inactivation varies in mouse tissues.","volume":"11","author":"Berletch","year":"2015","journal-title":"PLoS Genet."},{"key":"B13","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1007\/s00439-011-1011-z","article-title":"Genes that escape from X inactivation.","volume":"130","author":"Berletch","year":"2011","journal-title":"Hum. Genet."},{"key":"B14","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1038\/349038a0","article-title":"A gene from the region of the human X inactivation centre is expressed exclusively from the inactive X chromosome.","volume":"349","author":"Brown","year":"1991","journal-title":"Nature"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1186\/s13293-016-0115-5","article-title":"A primer on the use of mouse models for identifying direct sex chromosome effects that cause sex differences in non-gonadal tissues.","volume":"7","author":"Burgoyne","year":"2016","journal-title":"Biol. Sex Differ."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2016.0355","article-title":"When the Lyon(ized chromosome) roars: ongoing expression from an inactive X chromosome.","volume":"372","author":"Carrel","year":"2017","journal-title":"Philos. Trans. R. Soc. Lond. B. Biol. Sci."},{"key":"B17","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1038\/nature03479","article-title":"X-inactivation profile reveals extensive variability in X-linked gene expression in females.","volume":"434","author":"Carrel","year":"2005","journal-title":"Nature"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0023956","article-title":"The choice of the filtering method in microarrays affects the inference regarding dosage compensation of the active X-chromosome.","volume":"6","author":"Castagn\u00e9","year":"2011","journal-title":"PLoS One"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1259","DOI":"10.1101\/gr.075713.107","article-title":"DXZ4 chromatin adopts an opposing conformation to that of the surrounding chromosome and acquires a novel inactive X-specific role involving CTCF and antisense transcripts.","volume":"18","author":"Chadwick","year":"2008","journal-title":"Genome Res."},{"key":"B20","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1101\/gr.390503","article-title":"Complex events in the evolution of the human pseudoautosomal region 2 (PAR2).","volume":"13","author":"Charchar","year":"2003","journal-title":"Genome Res."},{"key":"B21","doi-asserted-by":"publisher","first-page":"2223","DOI":"10.1101\/gad.380906","article-title":"A novel role for Xist RNA in the formation of a repressive nuclear compartment into which genes are recruited when silenced.","volume":"20","author":"Chaumeil","year":"2006","journal-title":"Genes Dev."},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1038\/srep37324","article-title":"YY1 binding association with sex-biased transcription revealed through X-linked transcript levels and allelic binding analyses.","volume":"6","author":"Chen","year":"2016","journal-title":"Sci. Rep."},{"key":"B23","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1093\/hmg\/9.3.395","article-title":"Differentially regulated and evolved genes in the fully sequenced Xq\/Yq pseudoautosomal region.","volume":"9","author":"Ciccodicola","year":"2000","journal-title":"Hum. Mol. Genet."},{"key":"B24","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1083\/jcb.142.1.13","article-title":"Stabilization and localization of Xist RNA are controlled by separate mechanisms and are not sufficient for X inactivation.","volume":"142","author":"Clemson","year":"1998","journal-title":"J. Cell Biol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"7688","DOI":"10.1073\/pnas.0601069103","article-title":"The X chromosome is organized into a gene-rich outer rim and an internal core containing silenced nongenic sequences.","volume":"103","author":"Clemson","year":"2006","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B26","doi-asserted-by":"publisher","first-page":"488","DOI":"10.1038\/nature13151","article-title":"Origins and functional evolution of Y chromosomes across mammals.","volume":"508","author":"Cortez","year":"2014","journal-title":"Nature"},{"key":"B27","doi-asserted-by":"publisher","first-page":"1211","DOI":"10.1093\/hmg\/ddt513","article-title":"Spread of X-chromosome inactivation into autosomal sequences: role for DNA elements, chromatin features and chromosomal domains.","volume":"23","author":"Cotton","year":"2014","journal-title":"Hum. Mol. Genet."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1528","DOI":"10.1093\/hmg\/ddu564","article-title":"Landscape of DNA methylation on the X chromosome reflects CpG density, functional chromatin state and X-chromosome inactivation.","volume":"24","author":"Cotton","year":"2015","journal-title":"Hum. Mol. Genet."},{"key":"B29","doi-asserted-by":"publisher","first-page":"5491","DOI":"10.3168\/jds.2016-12490","article-title":"Bovine mammary gland X chromosome inactivation.","volume":"100","author":"Couldrey","year":"2017","journal-title":"J. Dairy Sci."},{"key":"B30","doi-asserted-by":"publisher","first-page":"E4504","DOI":"10.1073\/pnas.1609643113","article-title":"Deletion of DXZ4 on the human inactive X chromosome alters higher-order genome architecture.","volume":"113","author":"Darrow","year":"2016","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B31","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1007\/s10577-007-1121-6","article-title":"The search for a marsupial XIC reveals a break with vertebrate synteny.","volume":"15","author":"Davidow","year":"2007","journal-title":"Chromosome Res."},{"key":"B32","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1159\/000339433","article-title":"The evolution of marsupial and monotreme chromosomes.","volume":"137","author":"Deakin","year":"2012","journal-title":"Cytogenet. Genome Res."},{"key":"B33","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1038\/ng.948","article-title":"Evidence for compensatory upregulation of expressed X-linked genes in mammals, Caenorhabditis elegans and Drosophila melanogaster.","volume":"43","author":"Deng","year":"2011","journal-title":"Nat. Genet."},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-015-0728-8","article-title":"Bipartite structure of the inactive mouse X chromosome.","volume":"16","author":"Deng","year":"2015","journal-title":"Genome Biol."},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1004798","article-title":"Quantitative genetics of CTCF binding reveal local sequence effects and different modes of X-chromosome association.","volume":"10","author":"Ding","year":"2014","journal-title":"PLoS Genet."},{"key":"B36","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.semcdb.2016.04.013","article-title":"Dosage compensation of the sex chromosomes and autosomes.","volume":"56","author":"Disteche","year":"2016","journal-title":"Semin. Cell Dev. Biol."},{"key":"B37","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1038\/ng.3726","article-title":"Tumor-suppressor genes that escape from X-inactivation contribute to cancer sex bias.","volume":"49","author":"Dunford","year":"2017","journal-title":"Nat. Genet."},{"key":"B38","doi-asserted-by":"publisher","first-page":"1653","DOI":"10.1126\/science.1126316","article-title":"The Xist RNA gene evolved in eutherians by pseudogenization of a protein-coding gene.","volume":"312","author":"Duret","year":"2006","journal-title":"Science"},{"key":"B39","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1146\/annurev-genet-120116-024611","article-title":"X-chromosome inactivation: a crossroads between chromosome architecture and gene regulation.","volume":"52","author":"Galupa","year":"2018","journal-title":"Annu. Rev. Genet."},{"key":"B40","doi-asserted-by":"publisher","first-page":"13015","DOI":"10.1073\/pnas.1806811115","article-title":"Extensive cellular heterogeneity of X inactivation revealed by single-cell allele-specific expression in human fibroblasts.","volume":"115","author":"Garieri","year":"2018","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B41","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1146\/annurev.ge.17.120183.001103","article-title":"Mammalian X-chromosome inactivation.","volume":"17","author":"Gartler","year":"1983","journal-title":"Annu. Rev. Genet."},{"key":"B42","doi-asserted-by":"publisher","first-page":"575","DOI":"10.1038\/nature18589","article-title":"Structural organization of the inactive X chromosome in the mouse.","volume":"535","author":"Giorgetti","year":"2016","journal-title":"Nature"},{"key":"B43","doi-asserted-by":"publisher","first-page":"7542","DOI":"10.1073\/pnas.1505463112","article-title":"Evolutionary comparison reveals that diverging CTCF sites are signatures of ancestral topological associating domains borders.","volume":"112","author":"G\u00f3mez-Mar\u00edn","year":"2015","journal-title":"Proc. Natl. Acad. Sci. U.SA."},{"key":"B44","doi-asserted-by":"publisher","first-page":"638","DOI":"10.1002\/ca.22727","article-title":"Anatomy of turner syndrome: turner syndrome.","volume":"29","author":"Granger","year":"2016","journal-title":"Clin. Anat."},{"key":"B45","doi-asserted-by":"publisher","first-page":"254","DOI":"10.1038\/nature11171","article-title":"Rsx is a metatherian RNA with Xist-like properties in X-chromosome inactivation.","volume":"487","author":"Grant","year":"2012","journal-title":"Nature"},{"key":"B46","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1159\/000071585","article-title":"Evolution of the human X - a smart and sexy chromosome that controls speciation and development.","volume":"99","author":"Graves","year":"2002","journal-title":"Cytogenet. Genome Res."},{"key":"B47","doi-asserted-by":"publisher","first-page":"913","DOI":"10.1038\/nature03021","article-title":"In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes.","volume":"432","author":"Gr\u00fctzner","year":"2004","journal-title":"Nature"},{"key":"B48","doi-asserted-by":"crossref","DOI":"10.1186\/jbiol30","article-title":"Global analysis of X-chromosome dosage compensation.","author":"Gupta","year":"2006","journal-title":"J. Biol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"5133","DOI":"10.1073\/pnas.95.9.5133","article-title":"Reactivation of XIST in normal fibroblasts and a somatic cell hybrid: abnormal localization of XIST RNA in hybrid cells.","volume":"95","author":"Hansen","year":"1998","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B50","doi-asserted-by":"publisher","first-page":"2955","DOI":"10.1073\/pnas.1018392109","article-title":"A Y-linked anti-Mullerian hormone duplication takes over a critical role in sex determination.","volume":"109","author":"Hattori","year":"2012","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B51","doi-asserted-by":"publisher","first-page":"1171","DOI":"10.1038\/ng.1010","article-title":"He et al. Author reply.","volume":"43","author":"He","year":"2011","journal-title":"Nat. Genet."},{"key":"B52","doi-asserted-by":"publisher","first-page":"4367","DOI":"10.1093\/hmg\/dds270","article-title":"The macrosatellite DXZ4 mediates CTCF-dependent long-range intrachromosomal interactions on the human inactive X chromosome.","volume":"21","author":"Horakova","year":"2012","journal-title":"Hum. Mol. Genet."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1003952","article-title":"Deletion of an X-inactivation boundary disrupts adjacent gene silencing.","volume":"9","author":"Horvath","year":"2013","journal-title":"PLoS Genet."},{"key":"B54","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1038\/nature10843","article-title":"Strict evolutionary conservation followed rapid gene loss on human and rhesus Y chromosomes.","volume":"483","author":"Hughes","year":"2012","journal-title":"Nature"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1002315","article-title":"The constrained maximal expression level owing to haploidy shapes gene content on the mammalian X chromosome.","volume":"13","author":"Hurst","year":"2015","journal-title":"PLoS Biol."},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001328","article-title":"Mechanisms and evolutionary patterns of mammalian and avian dosage compensation.","volume":"10","author":"Julien","year":"2012","journal-title":"PLoS Biol."},{"key":"B57","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1016\/j.metabol.2017.09.017","article-title":"Klinefelter syndrome: more than hypogonadism.","volume":"86","author":"Kanakis","year":"2018","journal-title":"Metabolism"},{"key":"B58","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1002\/ajmg.1320420616","article-title":"Survival of fetuses with 45,X - an instructive case and an hypothesis.","volume":"42","author":"Kelly","year":"1992","journal-title":"Am. J. Med. Genet."},{"key":"B59","doi-asserted-by":"publisher","first-page":"1167","DOI":"10.1038\/ng.991","article-title":"Evidence for dosage compensation between the X chromosome and autosomes in mammals.","volume":"43","author":"Kharchenko","year":"2011","journal-title":"Nat. Genet."},{"key":"B60","doi-asserted-by":"publisher","first-page":"3964","DOI":"10.1093\/hmg\/ddr315","article-title":"Allele-specific distribution of RNA polymerase II on female X chromosomes.","volume":"20","author":"Kucera","year":"2011","journal-title":"Hum. Mol. Genet"},{"key":"B61","doi-asserted-by":"publisher","first-page":"964","DOI":"10.1126\/science.286.5441.964","article-title":"Four evolutionary strata on the human X chromosome.","volume":"286","author":"Lahn","year":"1999","journal-title":"Science"},{"key":"B62","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1093\/gbe\/evp006","article-title":"Footprints of inversions at present and past pseudoautosomal boundaries in human sex chromosomes.","volume":"1","author":"Lemaitre","year":"2009","journal-title":"Genome Biol. Evol."},{"key":"B63","doi-asserted-by":"publisher","first-page":"17055","DOI":"10.1073\/pnas.0807765105","article-title":"Escape from X chromosome inactivation is an intrinsic property of the Jarid1c locus.","volume":"105","author":"Li","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-017-00528-1","article-title":"Genetic and epigenetic features direct differential efficiency of Xist-mediated silencing at X-chromosomal and autosomal locations.","volume":"8","author":"Loda","year":"2017","journal-title":"Nat. Commun."},{"key":"B65","doi-asserted-by":"publisher","first-page":"2656","DOI":"10.1128\/MCB.00183-16","article-title":"Xist and Tsix transcription dynamics is regulated by the X-to-autosome ratio and semistable transcriptional states.","volume":"36","author":"Loos","year":"2016","journal-title":"Mol. Cell. Biol."},{"key":"B66","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1038\/190372a0","article-title":"Gene action in the X of the mouse.pdf.","volume":"190","author":"Lyon","year":"1961","journal-title":"Nature"},{"key":"B67","first-page":"135","article-title":"Sex chromatin and gene action in the mammalian X-chromosome.","volume":"14","author":"Lyon","year":"1962","journal-title":"Am. J. Hum. Genet."},{"key":"B68","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1159\/000014969","article-title":"X-chromosome inactivation: a repeat hypothesis.","volume":"80","author":"Lyon","year":"1998","journal-title":"Cytogenet. Genome Res."},{"key":"B69","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1177\/1098612X15571878","article-title":"DNA mutations of the cat: the good, the bad and the ugly.","volume":"17","author":"Lyons","year":"2015","journal-title":"J. Feline Med. Surg."},{"key":"B70","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1016\/j.tig.2013.07.005","article-title":"Sex chromosome dosage compensation: definitely not for everyone.","volume":"29","author":"Mank","year":"2013","journal-title":"Trends Genet."},{"key":"B71","doi-asserted-by":"publisher","first-page":"2133","DOI":"10.1111\/j.1558-5646.2011.01316.x","article-title":"Some inconvenient truths about sex chromosome dosage compensation and the potential role of sexual conflict.","volume":"65","author":"Mank","year":"2011","journal-title":"Evolution"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-015-0698-x","article-title":"Dynamics of gene silencing during X inactivation using allele-specific RNA-seq.","volume":"16","author":"Marks","year":"2015","journal-title":"Genome Biol."},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0170403","article-title":"Embryonic loss of human females with partial trisomy 19 identifies region critical for the single active X.","volume":"12","author":"Migeon","year":"2017","journal-title":"PLoS One"},{"key":"B74","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1534\/genetics.114.162800","article-title":"Evidence for local regulatory control of escape from imprinted X chromosome inactivation.","volume":"197","author":"Mugford","year":"2014","journal-title":"Genetics"},{"key":"B75","doi-asserted-by":"publisher","first-page":"350","DOI":"10.1038\/s41594-019-0214-1","article-title":"A symmetric toggle switch explains the onset of random X inactivation in different mammals.","volume":"26","author":"Mutzel","year":"2019","journal-title":"Nat. Struct. 38 Mol. Biol."},{"key":"B76","doi-asserted-by":"publisher","first-page":"474","DOI":"10.1101\/gr.230433.117","article-title":"Conserved microRNA targeting reveals preexisting gene dosage sensitivities that shaped amniote sex chromosome evolution.","volume":"28","author":"Naqvi","year":"2018","journal-title":"Genome Res."},{"key":"B77","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1016\/j.tig.2019.04.008","article-title":"The pregnancy pickle: evolved immune compensation due to pregnancy underlies sex differences in human diseases.","volume":"35","author":"Natri","year":"2019","journal-title":"Trends Genet."},{"key":"B78","doi-asserted-by":"publisher","first-page":"1375","DOI":"10.1002\/ajmg.a.38710","article-title":"X-linked intellectual disability update 2017.","volume":"176","author":"Neri","year":"2018","journal-title":"Am. J. Med. Genet. B Neuropsychiatr. Genet."},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0113434","article-title":"Roles of intragenic and intergenic L1s in mouse and human.","volume":"9","author":"Ngamphiw","year":"2014","journal-title":"PLoS One"},{"key":"B80","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1038\/ng1705","article-title":"Dosage compensation of the active X chromosome in mammals.","volume":"38","author":"Nguyen","year":"2006","journal-title":"Nat. Genet."},{"key":"B81","author":"Nussbaum","year":"2001","journal-title":"Thompson & Thompson Genetics in Medicine"},{"key":"B82","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-642-88178-7","author":"Ohno","year":"1967","journal-title":"Sex Chromosomes and Sex-Linked Genes."},{"key":"B83","doi-asserted-by":"publisher","first-page":"1863","DOI":"10.1093\/gbe\/evt139","article-title":"Detecting evolutionary strata on the human X chromosome in the absence of gametologous Y-linked sequences.","volume":"5","author":"Pandey","year":"2013","journal-title":"Genome Biol. Evol."},{"key":"B84","doi-asserted-by":"publisher","first-page":"1252","DOI":"10.1093\/hmg\/ddy039","article-title":"Human cis-acting elements regulating escape from X-chromosome inactivation function in mouse.","volume":"27","author":"Peeters","year":"2018","journal-title":"Hum. Mol. Genet."},{"key":"B85","doi-asserted-by":"publisher","first-page":"1383","DOI":"10.1007\/s00018-013-1499-6","article-title":"The evolution of X chromosome inactivation in mammals: the demise of Ohno\u2019s hypothesis?","volume":"71","author":"Pessia","year":"2014","journal-title":"Cell. Mol. Life Sci."},{"key":"B86","doi-asserted-by":"publisher","first-page":"5346","DOI":"10.1073\/pnas.1116763109","article-title":"Mammalian X chromosome inactivation evolved as a dosage-compensation mechanism for dosage-sensitive genes on the X chromosome.","volume":"109","author":"Pessia","year":"2012","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B87","doi-asserted-by":"publisher","first-page":"492","DOI":"10.1016\/j.tig.2018.04.001","article-title":"Why do sex chromosomes stop recombining?","volume":"34","author":"Ponnikas","year":"2018","journal-title":"Trends Genet."},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.0060080","article-title":"Chromosomal gene movements reflect the recent origin and biology of therian sex chromosomes.","volume":"6","author":"Potrzebowski","year":"2008","journal-title":"PLoS Biol."},{"key":"B89","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/j.cels.2015.06.003","article-title":"Individuality and variation of personal regulomes in primary human T cells.","volume":"1","author":"Qu","year":"2015","journal-title":"Cell Syst."},{"key":"B90","doi-asserted-by":"publisher","first-page":"1665","DOI":"10.1016\/j.cell.2014.11.021","article-title":"A 3D map of the human genome at kilobase resolution reveals principles of chromatin looping.","volume":"159","author":"Rao","year":"2014","journal-title":"Cell"},{"key":"B91","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1159\/000330627","article-title":"The pseudoautosomal region and sex chromosome aneuploidies in domestic species.","volume":"6","author":"Raudsepp","year":"2012","journal-title":"Sex. Dev."},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2007-8-11-r243","article-title":"The multiple sex chromosomes of platypus and echidna are not completely identical and several share homology with the avian Z.","volume":"8","author":"Rens","year":"2007","journal-title":"Genome Biol."},{"key":"B93","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1111\/j.1558-5646.1984.tb00346.x","article-title":"Sex chromosomes and the evolution of sexual dimorphism.","volume":"38","author":"Rice","year":"1984","journal-title":"Evolution"},{"key":"B94","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1038\/newbio230154a0","article-title":"Inheritance of glucose-6-phosphate dehydrogenase variation in kangaroos.","volume":"230","author":"Richardson","year":"1971","journal-title":"Nature. New Biol."},{"key":"B95","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1038\/nature03440","article-title":"The DNA sequence of the human X chromosome.","volume":"434","author":"Ross","year":"2005","journal-title":"Nature"},{"key":"B96","doi-asserted-by":"crossref","first-page":"969","DOI":"10.1242\/dev.110.3.969","article-title":"An extra maternally derived X chromosome is deleterious to early mouse development.","volume":"110","author":"Shao","year":"1990","journal-title":"Development"},{"key":"B97","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1038\/346240a0","article-title":"A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif.","volume":"346","author":"Sinclair","year":"1990","journal-title":"Nature"},{"key":"B98","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1038\/nature01722","article-title":"The male-specific region of the human Y chromosome is a mosaic of discrete sequence classes.","volume":"423","author":"Skaletsky","year":"2003","journal-title":"Nature"},{"key":"B99","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1038\/nature08298","article-title":"The avian Z-linked gene DMRT1 is required for male sex determination in the chicken.","volume":"461","author":"Smith","year":"2009","journal-title":"Nature"},{"key":"B100","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1016\/0888-7543(91)90141-Z","article-title":"Genes on the short arm of the human X chromosome are not shared with the marsupial X.","volume":"11","author":"Spencer","year":"1991","journal-title":"Genomics"},{"key":"B101","doi-asserted-by":"publisher","first-page":"2013","DOI":"10.1093\/hmg\/2.12.2013","article-title":"SOX3 is an X-linked gene related to SRY.","volume":"2","author":"Stevanovl\u0107","year":"1993","journal-title":"Hum. Mol. Genet."},{"key":"B102","doi-asserted-by":"publisher","first-page":"640","DOI":"10.1038\/256640a0","article-title":"Preferential inactivation of the paternally derived X chromosome in the extraembryonic membranes of the mouse.","volume":"256","author":"Takagi","year":"1975","journal-title":"Nature"},{"key":"B103","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1038\/nature24265","article-title":"Landscape of X chromosome inactivation across human tissues.","volume":"550","author":"Tukiainen","year":"2017","journal-title":"Nature"},{"key":"B104","doi-asserted-by":"publisher","first-page":"965","DOI":"10.1101\/gr.7101908","article-title":"Bird-like sex chromosomes of platypus imply recent origin of mammal sex chromosomes.","volume":"18","author":"Veyrunes","year":"2008","journal-title":"Genome Res."},{"key":"B105","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1101\/gr.161919.113","article-title":"Chromosome-wide profiling of X-chromosome inactivation and epigenetic states in fetal brain and placenta of the opossum.","volume":"24","author":"Wang","year":"2014","journal-title":"Monodelphis domestica. Genome Res."},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.0020113","article-title":"Evidence of influence of genomic DNA sequence on human X chromosome inactivation.","volume":"2","author":"Wang","year":"2006","journal-title":"PLoS Comput. Biol."},{"key":"B107","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1016\/j.molcel.2012.11.013","article-title":"Latent regulatory potential of human-specific repetitive elements.","volume":"49","author":"Ward","year":"2013","journal-title":"Mol. Cell."},{"key":"B108","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1093\/molbev\/msx297","article-title":"Landscape of DNA methylation on the marsupial X.","volume":"35","author":"Waters","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"B109","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1016\/j.semcdb.2016.01.006","article-title":"The X factor: X chromosome dosage compensation in the evolutionarily divergent monotremes and marsupials.","volume":"56","author":"Whitworth","year":"2016","journal-title":"Semin. Cell Dev. Biol."},{"key":"B110","doi-asserted-by":"publisher","first-page":"1043","DOI":"10.1038\/ng.711","article-title":"RNA sequencing shows no dosage compensation of the active X-chromosome.","volume":"42","author":"Xiong","year":"2010","journal-title":"Nat. Genet."},{"key":"B111","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/j.ygeno.2007.07.002","article-title":"A cross-species comparison of X-chromosome inactivation in Eutheria.","volume":"90","author":"Yen","year":"2007","journal-title":"Genomics"}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fcell.2019.00241\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,10,2]],"date-time":"2022-10-02T08:10:16Z","timestamp":1664698216000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fcell.2019.00241\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,10,22]]},"references-count":111,"alternative-id":["10.3389\/fcell.2019.00241"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2019.00241","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,10,22]]},"article-number":"241"}}