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. 2019 Dec 17;9:19232. doi: 10.1038/s41598-019-55291-8

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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DMRs distally located with respect to genes are enriched in biologically relevant pathways. (A) Pathways analysis using the Genomic Regions Enrichment of Annotations Tool (GREAT) that associates each gene with a ‘regulatory domain’ defined as a genomic region 5 kb upstream and 1 kb downstream from the TSS and an extension within 1 Mb up to the regulatory domain of the nearest upstream or downstream gene. (B) Schematic model depicting methylation-dependent modulation of TF binding to distal regulatory domains. Left diagram, hypermethylation of an activator inhibits, while hypomethylation of an activator facilitates TF binding, resulting in transcriptional repression or activation, respectively. Right diagram, particular TFs bind hypermethylated repressors, leading to transcriptional repression, while hypomethylation has the opposite effect on transcription. Bottom diagram, TF binding to a regulatory domain enables long-distance interactions with the target genes through 3D chromatin structural rearrangement.