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First published online July 22, 2005; 10.1104/pp.105.063537

Plant Physiology 138:1842-1852 (2005)
© 2005 American Society of Plant Biologists

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An RNA-Dependent RNA Polymerase Prevents Meristem Invasion by Potato Virus X and Is Required for the Activity But Not the Production of a Systemic Silencing Signal1,[w]

Frank Schwach, Fabian E. Vaistij2, Louise Jones2 and David C. Baulcombe*

Sainsbury Laboratory, Norwich NR4 7UH, United Kingdom

One of the functions of RNA silencing in plants is antiviral defense. A hallmark of RNA silencing is spreading of the silenced state through the plant. Little is known about the nature of the systemic silencing signal and the proteins required for its production, transport, and reception in plant tissues. Here, we show that the RNA-dependent RNA polymerase RDR6 in Nicotiana benthamiana is involved in defense against potato virus X at the level of systemic spreading and in exclusion of the virus from the apical growing point. It has no effect on primary replication and cell-to-cell movement of the virus and does not contribute significantly to the formation of virus-derived small interfering (si) RNA in a fully established potato virus X infection. In grafting experiments, the RDR6 homolog was required for the ability of a cell to respond to, but not to produce or translocate, the systemic silencing signal. Taking these findings together, we suggest a model of virus defense in which RDR6 uses incoming silencing signal to generate double-stranded RNA precursors of secondary siRNA. According to this idea, the secondary siRNAs mediate RNA silencing as an immediate response that slows down the systemic spreading of the virus into the growing point and newly emerging leaves.

1 This work was supported by the European Commission Research Directorate (contract no. QLG2–CT–2002–01673). The Sainsbury Laboratory is supported by the Gatsby Charitable Foundation.

2 Present address: Department of Biology, University of York, York YO10 5DD, UK.

[w] The online version of this article contains Web-only data.

Article, publication date, and citation information can be found at www.plantphysiol.org/cgi/doi/10.1104/pp.105.063537.

* Corresponding author; e-mail david.baulcombe{at}sainsbury-laboratory.ac.uk; fax (44)(0)1603–450011.

Received March 30, 2005; returned for revision April 20, 2005; accepted April 20, 2005.




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