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Our clustering analysis reveals that considerable diversity of nucleosome position, occupancy, and remodeling underlies the central mode of remodeling.
Prior to GR binding, the enhancers belonging to each mode of remodeling also have very different properties.
For genes associated with multiple proximal GR peaks, we considered only those genes for which all peaks have the same mode of remodeling.
Finally, we used the Genomic Regions Enrichment of Annotations Tool (GREAT, [ 30]) to apply gene expression-based gene ontologies in a "pathway" analysis of GR binding sites for each mode of remodeling.
He et al. investigated nucleosome dynamics relative to chromatin binding by androgen receptor and used H3K4me2 ChIP-Seq to describe nucleosome depletion overlapping transcription factor binding that is similar to the "central" mode of remodeling we define here [ 12].
In aggregate, we see diminished central ChIP-Seq signal without corresponding increases in signal of flanking nucleosomes, suggesting that nucleosome eviction, rather than repositioning, predominates within this central mode of remodeling.
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Both measures of enhancer activation demonstrate significant differences between modes of remodeling.
Based upon these patterns, we grouped the GR binding sites into four "modes" of remodeling.
The remaining regions of minimal remodeling also have low levels of enhancer activation, bearing less H3K27Ac and DHS relative to other modes of remodeling (Fig. 5b, c).
Unlike other modes of remodeling, the total H3K4methyls signal at sites of phased remodeling increases, suggesting recruitment of histone methyltransferases or displacement of histone demethylases (data not shown).
We first evaluated the GR binding and enhancer properties of sites belonging to each mode of nucleosome remodeling.
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