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Molecular and Cellular Biology, October 2008, p. 6452-6461, Vol. 28, No. 20
0270-7306/08/$08.00+0 doi:10.1128/MCB.01015-08
Copyright © 2008, American Society for Microbiology. All Rights Reserved.

Xian Zhang,
William D. Burke, and
Thomas H. Eickbush*
Department of Biology, University of Rochester, Rochester, New York 14627
Received 27 June 2008/ Returned for modification 22 July 2008/ Accepted 29 July 2008
R2 retrotransposable elements exclusively insert into a conserved region of the tandemly organized 28S rRNA genes. Despite inactivating a subset of these genes, R2 elements have persisted in the ribosomal DNA (rDNA) loci of insects for hundreds of millions of years. Controlling R2 proliferation was addressed in this study using lines of Drosophila simulans previously shown to have either active or inactive R2 retrotransposition. Lines with active retrotransposition were shown to have high R2 transcript levels, which nuclear run-on transcription experiments revealed were due to increased transcription of R2-inserted genes. Crosses between R2 active and inactive lines indicated that an important component of this transcriptional control is linked to or near the rDNA locus, with the R2 transcription level of the inactive parent being dominant. Pulsed-field gel analysis suggested that the R2 active and inactive states were determined by R2 distribution within the locus. Molecular and cytological analyses further suggested that the entire rDNA locus from the active line can be silenced in favor of the locus from the inactive line. This silencing of entire rDNA loci represents an example of the large-scale epigenetic control of transposable elements and shares features with the nucleolar dominance frequently seen in interspecies hybrids.
Published ahead of print on 4 August 2008.
Present address: Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA.
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