6.4: Contact-dependent Signaling
Contact-dependent signaling uses specialized cytoplasmic channels between cells that allow the flow of small molecules between them. In animal cells, these channels are called gap junctions. In plants, they are known as plasmodesmata.
Gap junctions form when two hemichannels, or connexons, join; one connexon from one cell coupling to a connexon of an adjacent cell. Each cell’s connexon is formed from six proteins creating a circular channel. There are over 20 different types of these proteins, or connexins, so there is substantial variation in how they come together as connexons and as gap junctions. Connexins have four transmembrane subunits with both their N- and C-terminus endings located intracellularly. The C-terminus has multiple phosphorylation sites so it can be activated by numerous different kinases- further adding to gap junction variety.
Depending on the activating kinase, and the C-terminal amino acid residues of connexins that are phosphorylated, gap junctions can be partially or fully opened. This selectively allows small molecules to flow from one cell into another. A gap junction may also exclude by electrochemical charge. The selectivity of gap junctions allows a single cell to coordinate a complex multicellular response. However, some toxic molecules, matching the size and electrochemical preference of the gap junction, can also pass between cells. This results in a “bystander effect.” where a damaged or diseased cell passes on apoptotic signals to its neighbor resulting in the death of both the cells.
In plants, contact-dependent signaling occurs through the use of plasmodesmata. Like gap junctions, they are cytoplasmic channels between two cells. They pass through both the cell wall and membrane. The plasmodesmata also direct communication from a single cell out to many cells; thus they are also susceptible to “bystander effects.”
Unlike gap junctions, the channels of plasmodesmata are more flexible in the molecules they let pass between cells. At the center of most channels is a tube of the endoplasmic reticulum (ER) that runs between the two cells. This ER extension is called the desmotubule which surrounds the cytoplasmic sleeve. This sleeve permits the passive and active transport of molecules. To transport larger molecules, like transcription factors, small RNA, or other nucleic acids and proteins, the plasmodesmata can dilate. Because of this ability, plasmodesmata play a key role in cell position information and cell fate. Unfortunately, this ability also permits viral infection to quickly spread between cells.