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doi:10.1534/genetics.106.058669
A more recent version of this article appeared on July 1, 2006.
REGULAR RESEARCH PAPERS |
Transcriptionally Silenced Transgenes in Maize are Activated by Three Mutations Defective in Paramutation
Karen M. McGinnis 1, Catherine Springer 2, Yan Lin 1, Charles C. Carey 3 and Vicki L. Chandler 1*
1 University of Arizona
2 University of Minnesota
3 Fred Hutchinson Cancer Research Center
* To whom correspondence should be addressed. E-mail: chandler{at}ag.arizona.edu.
Submitted on March 28, 2006
Revised on April 11, 2006
Accepted on 9 May 2006
Plants with mutations in one of three maize genes, mop1, rmr1 and rmr2, are defective in paramutation, an allele-specific interaction that leads to meiotically heritable chromatin changes. Experiments reported here demonstrate these genes are required to maintain the transcriptional silencing of two different transgenes, suggesting paramutation and transcriptional silencing of transgenes share mechanisms. We hypothesize the transgenes are silenced through an RNA-directed chromatin mechanism, because mop1 encodes an RNA-dependent RNA polymerase. In all the mutants, DNA methylation was reduced in the active transgenes relative to the silent transgenes at all of the CNG sites monitored within the transgene promoter. However, asymmetrical methylation persisted at one site within the reactivated transgene in the rmr1-1 mutant. With that one mutant, rmr1-1, the transgene was efficiently resilenced upon outcrossing to reintroduce the wild type protein. In contrast, with the mop1-1 and rmr2-1 mutants, the transgene remained active in a subset of progeny even after the wild type proteins were reintroduced by outcrossing. Interestingly, this immunity to silencing increased as the generations progressed, consistent with a heritable chromatin state being formed at the transgene in plants carrying the mop1-1 and rmr2-1 mutations that becomes more resistant to silencing in subsequent generations.
Key Words: RNA dependent RNA polymerase, RNA directed chromatin structure, paramutation, transcriptional silencing, transgenes
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