The plasmodesmatal transport pathway for homoeotic proteins, silencing signals and viruses

Non-cell-autonomous signals in the form of microRNAs and transcription factors could have important developmental functions. Plasmodesmata (PD) form a cytoplasmic network throughout the plant body and provide the means of symplasmic cell-to-cell transport in plants. Homeodomain transcription factors, small RNA molecules and viral genomic information move selectively to adjacent cells via PD microchannels. Tissue-specific expression studies of non-cell-autonomous transcription factors and RNA molecules have confirmed that their intercellular transport is a highly regulated process, which depends on the tissue, developmental stage and nature of the transported macromolecule. We have known for some time that gene-silencing signals spread both locally from cell to cell and across long distances following the source to sink transition. Recent work has provided evidence that small single-stranded silencing-induced RNAs and microRNA molecules are present in the phloem transport system of different plant species. Further, recent evidence has confirmed that the transport of silencing RNA via PD is a regulated and active process, and that an amplification-relay mechanism is in place for the long-distance spread of silencing signals.