Key checkpoints in the movement of plant viruses through the host.

Plant viruses cannot exploit any of the membrane fusion-based routes of entry described for animal viruses. In addition, one of the distinctive structures of plant cells, the cell wall, acts as the first barrier against the invasion of pathogens. To overcome the rigidity of the cell wall, plant viruses normally take advantage of the way of life of different biological vectors. Alternatively, the physical damage caused by environmental stresses can facilitate virus entry. Once inside the cell and taking advantage of the characteristic symplastic continuity of plant cells, viruses need to remodel and/or modify the restricted pore size of the plasmodesmata (channels that connect plant cells). In a successful interaction for the virus, it can reach the vascular tissue to systematically invade the plant. The connections between the different cell types in this path are not designed to allow the passage of molecules with the complexity of viruses. During this process, viruses face different cell barriers that must be overcome to reach the distal parts of the plant. In this review, we highlight the current knowledge about how plant RNA viruses enter plant cells, move between them to reach vascular cells and overcome the different physical and cellular barriers that the phloem imposes. Finally, we update the current research on cellular organelles as key regulator checkpoints in the long-distance movement of plant viruses.