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Transcription and RNA interference in the formation of heterochromatin

TLDR
Transcription in heterochromatin seems to be an oxymoron — surely the 'silenced' form of chromatin should not be transcribed, but there have been frequent reports of low-level transcription inheterochromatic regions, and several hundred genes are found in these regions in Drosophila.
Abstract
Transcription in heterochromatin seems to be an oxymoron--surely the 'silenced' form of chromatin should not be transcribed. But there have been frequent reports of low-level transcription in heterochromatic regions, and several hundred genes are found in these regions in Drosophila. Most strikingly, recent investigations implicate RNA interference mechanisms in targeting and maintaining heterochromatin, and these mechanisms are inherently dependent on transcription. Silencing of chromatin might involve trans-acting sources of the crucial small RNAs that carry out RNA interference, but in some cases, transcription of the region to be silenced seems to be required--an apparent contradiction.

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Control of the Embryonic Stem Cell State

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The Piwi-piRNA Pathway Provides an Adaptive Defense in the Transposon Arms Race

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Histones: annotating chromatin.

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References
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Journal ArticleDOI

Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans

TL;DR: To their surprise, it was found that double-stranded RNA was substantially more effective at producing interference than was either strand individually, arguing against stochiometric interference with endogenous mRNA and suggesting that there could be a catalytic or amplification component in the interference process.
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Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain.

TL;DR: A stepwise model for the formation of a transcriptionally silent heterochromatin is provided: SUV39H1 places a ‘methyl marker’ on histone H3, which is then recognized by HP1 through its chromo domain, which may also explain the stable inheritance of theheterochromatic state.
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Regulation of chromatin structure by site-specific histone H3 methyltransferases

TL;DR: A functional interdependence of site-specific H3 tail modifications is revealed and a dynamic mechanism for the regulation of higher-order chromatin is suggested.
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Discrete Small RNA-Generating Loci as Master Regulators of Transposon Activity in Drosophila

TL;DR: Examination of piwi-interacting RNAs associated with each Drosophila Piwi protein finds that Piwi and Aubergine bind RNAs that are predominantly antisense to transposons, whereas Ago3 complexes contain predominantly sense piRNAs.
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Regulation of heterochromatic silencing and histone H3 lysine-9 methylation by RNAi.

TL;DR: It is proposed that double-stranded RNA arising from centromeric repeats targets formation and maintenance of heterochromatin through RNAi.
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