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      The evolution of RNAi as a defence against viruses and transposable elements

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          Abstract

          RNA interference (RNAi) is an important defence against viruses and transposable elements (TEs). RNAi not only protects against viruses by degrading viral RNA, but hosts and viruses can also use RNAi to manipulate each other's gene expression, and hosts can encode microRNAs that target viral sequences. In response, viruses have evolved a myriad of adaptations to suppress and evade RNAi. RNAi can also protect cells against TEs, both by degrading TE transcripts and by preventing TE expression through heterochromatin formation. The aim of our review is to summarize and evaluate the current data on the evolution of these RNAi defence mechanisms. To this end, we also extend a previous analysis of the evolution of genes of the RNAi pathways. Strikingly, we find that antiviral RNAi genes, anti-TE RNAi genes and viral suppressors of RNAi all evolve rapidly, suggestive of an evolutionary arms race between hosts and parasites. Over longer time scales, key RNAi genes are repeatedly duplicated or lost across the metazoan phylogeny, with important implications for RNAi as an immune defence.

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          Most cited references134

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          Control of translation and mRNA degradation by miRNAs and siRNAs.

          The control of translation and mRNA degradation is an important part of the regulation of gene expression. It is now clear that small RNA molecules are common and effective modulators of gene expression in many eukaryotic cells. These small RNAs that control gene expression can be either endogenous or exogenous micro RNAs (miRNAs) and short interfering RNAs (siRNAs) and can affect mRNA degradation and translation, as well as chromatin structure, thereby having impacts on transcription rates. In this review, we discuss possible mechanisms by which miRNAs control translation and mRNA degradation. An emerging theme is that miRNAs, and siRNAs to some extent, target mRNAs to the general eukaryotic machinery for mRNA degradation and translation control.
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            Mechanisms of gene silencing by double-stranded RNA.

            Double-stranded RNA (dsRNA) is an important regulator of gene expression in many eukaryotes. It triggers different types of gene silencing that are collectively referred to as RNA silencing or RNA interference. A key step in known silencing pathways is the processing of dsRNAs into short RNA duplexes of characteristic size and structure. These short dsRNAs guide RNA silencing by specific and distinct mechanisms. Many components of the RNA silencing machinery still need to be identified and characterized, but a more complete understanding of the process is imminent.
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              RNA silencing in plants.

              There are at least three RNA silencing pathways for silencing specific genes in plants. In these pathways, silencing signals can be amplified and transmitted between cells, and may even be self-regulated by feedback mechanisms. Diverse biological roles of these pathways have been established, including defence against viruses, regulation of gene expression and the condensation of chromatin into heterochromatin. We are now in a good position to investigate the full extent of this functional diversity in genetic and epigenetic mechanisms of genome control.
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                Author and article information

                Journal
                Philos Trans R Soc Lond B Biol Sci
                RSTB
                Philosophical Transactions of the Royal Society B: Biological Sciences
                The Royal Society (London )
                0962-8436
                1471-2970
                16 October 2008
                12 January 2009
                : 364
                : 1513 , Theme Issue ‘Ecological immunology’ compiled by Hinrich Schulenburg, Joachim Kurtz, Yannick Moret and Mike T. Siva-Jothy
                : 99-115
                Affiliations
                [1 ]Institute of Evolutionary Biology, School of Biological Sciences, The University of Edinburgh, Ashworth Laboratories The King's Buildings, Edinburgh EH9 3JT, UK
                [2 ]CSIRO Entomology GPO Box 1700, Canberra ACT 2601, Australia
                [3 ]Division of Pathway Medicine and Centre for Infectious Diseases, University of Edinburgh Medical School Chancellor's Building, 49 Little France Crescent, Edinburgh EH16 4SB, UK
                Author notes
                [* ]Author for correspondence ( darren.obbard@ 123456ed.ac.uk )
                Article
                rstb20080168
                10.1098/rstb.2008.0168
                2592633
                18926973
                43c3c1d9-853c-4f7b-8271-08f1e5a53bf9
                Copyright © 2008 The Royal Society

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                Categories
                Review

                Philosophy of science
                viral suppressors of rnai,mirna,host–parasite coevolution,transposable element,pirna,antiviral rnai

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