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A series of recent studies have revealed that mouse and human ES cells possess certain novel epigenetic features. Polycomb-group (PcG) complex proteins mainly act to stabilize a repressive chromatin structure. Polycomb repressive complex 2 (PRC2), which consists of Ezh2, Eed and Suz12 in ES cells, functions as a histone methyltransferase on lysine 27 (K27) of histone H3, resulting in its tri-methylation (H3K27me3), a methylation mark that is associated with transcriptionally inactive genes ( Furtrimmed bomber jacket Yves Salomon Army Buy Cheap Free Shipping Discount For Cheap Authentic Free Shipping Discounts Sale Get Authentic Xwxlw
). In general, the distribution of this repressive chromatin mark is mutually exclusive to that of the tri-methylation mark H3K4me3, which is associated with transcriptionally active regions ( Strahl and Allis, 2000 ; Babs Skinny Cropped Jeans T Tahari Free Shipping Browse JrkOYP3qkv
). However, Bernstein et al. reported that in mouse ES cells, these histone marks co-localize in particular regions, which they named `bivalent domains' ( Pay With Paypal Free Shipping Classic Prada Tweed Virgin Wool amp; LinenBlend Jacket Marketable For Sale MM7X8KHO
). These domains, which are composed of short chromatin elements marked by H3K4me3 flanked by larger regions that contain H3K27me3, are associated with genes that are expressed at low levels ( Fig. 5B ) ( Professional Discount Online Hache Virgin Wool KneeLength Dress Mk7OL1
). Interestingly, the bivalent domains map to highly conserved noncoding elements (HCNEs) that have previously been identified as being conserved among the genomes of primates and rodents and which contain few retrotransposons ( Bernstein et al., 2006 ). Moreover, half of these bivalent domains contain target sites that are common to Oct3/4, Sox2 and Nanog, as identified by genome-wide ChIP-on-Chip analysis ( Boyer et al., 2005 ). Thus, these domains might signify the chromatin structure of genes that are in a differentiation-ready state, as proposed in the `Localised Marking Model' by Szutoristz and Dillon ( Szutoristz and Dillon, 2005 ). According to this model, most tissue-specific genes in ES cells would be targets for sequence-specific factors that can recruit histone-modifying enzymes, resulting in the formation of early transcription competence marks (ETCMs), which are enriched for histone H3 and H4 acetylation (H3Ac and H4Ac, respectively), and H3K4me3, all of which are histone marks associated with transcriptionally active regions. In both bivalent domains and ETCMs, H3K4me3 marks spread as genes near them become transcriptionally active, whereas H3K27me3 exclusively occupies those genes that are repressed during the differentiation of a particular cell type. Because the global level of H3K27me3 in ES cells is lower than that in differentiated cells, the mechanism by which this repressive mark targets such sites is of interest. Lee et al. ( Lee et al., 2006 ) performed ChIP-on-Chip analysis for Suz12, Eed and H3K27me3, and revealed that Suz12- and Eed-binding sites significantly overlap with each other and with H3K27me3 marks on the highly evolutionarily-conserved regions of transcriptionally silent genes, including Gata4 and Cdx2 , in ES cells. The 1800 genes identified as targets of Suz12 included most of the targets repressed by Oct3/4, Sox2 and Nanog ( Boyer et al., 2005 ). Boyer et al. ( Cheap Eastbay Discount High Quality Rag amp; Bone MidRise Silk Shorts Visa Payment Cheap Online Pick A Best Sale Online LBt1Rb
) also identified 512 common target genes of PRC2 and PRC1 by ChIP-on-Chip analysis and found that they were marked by H3K27me3, and that 87% were upregulated in the absence of PRC2 in Eed -null ES cells.

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