Plants and animals have had about 1.6 billion years-the time that has passed since they diverged from their last unicellular ancestor-to evolve different mechanisms for solving unique problems of development and intercellular signaling. As an example, plant cells are separated from each other by a substantial extracellular matrix, the cell wall. Although the cell wall is not impervious, it is likely to hinder cell-to-cell communication. Moreover, because plant cells cannot migrate during development, location, instead of lineage, is the primary determinant of cell fate in plants, making communication over both long and short distances essential for the coordination of plant growth. It appears that plants have significantly overcome the downside of having a cell wall by forming channels between cells to allow the transit of signaling and other important molecules. Plants may even be considered as supracellular organisms, in that whole tissues are symplastically connected. The channels that connect plant cells are called plasmodesmata (PD), and their investigation offers tantalizing clues as to how plant cells may use them to communicate with each other and to coordinate development. Although once seen as constricted channels through which only molecules <1 kD in size could pass (Terry and Robards 1987;Burnell 1988), PD have recently been shown to be far more dynamic and allow passage of both proteins and nucleic acids. Other findings, such as the observation that transcription factors can move between cells, and that RNA movement may be behind a range of long-distance signals in plants, raise further questions regarding a regulatory role of PD in development. Because of the importance of PD, we first summarize what we know about them, and then go on to discuss intercellular movement of proteins and RNA in general.
Plasmodesmata
UltrastructureUltrastructural studies of the PD by transmission electron microscopy (TEM) have shown them to be membrane-lined pores, with an exterior membrane contributed by the plasma membranes of the two cells connected by the PD, and a central desmotubule, derived from endoplasmic reticulum that is also contiguous between the two cells (Ding 1998). Typically, PD have a small diameter of ∼40 nm (Robards and Lucas 1990; van Bel and van Kesteren 1999), with the central desmotubule being connected to the plasma membrane via spoke-like extensions, resulting in microchannels having an estimated diameter of ∼2.5 nm (see Fig. 1). It is important to bear in mind that these measurements are all based on fixed tissue, and that nothing is known from these observations about the dynamic behavior of PD proteins.Morphologically, two distinct forms of PD have been recognized. Primary PD are formed at the cell plate, a plant-typical structure that consists of the incipient plasma membrane and cell wall synthesized during cytokinesis. Secondary PD, in contrast, are formed between existing cell walls. Both forms of PD can be either single-channels or branched. It is not known whether these different form...