When a cell in an intact plant growing in soil loses water, turgor declines and solute concentrations increase. As explained before (3.1.1), at turgor loss point, when turgor becomes zero, the hydrostatic pressure in the cell sap is equal to the atmospheric pressure, meaning that no net force is exerted on the cell wall, and the plant is wilting. If water continues to be lost from the cell, the pressure within the cytoplasm drops below atmospheric pressure, resulting in a force imbalance that collapses the cell wall. The deformation of living cells upon desiccation is called cytorrhysis. Note that the plasma membrane remains in close contact with the cell wall throughout desiccation ie plasmolysis does not occur, because the hydrostatic pressure in the cytoplasm remains greater than the hydrostatic pressure in the apoplast.
Plasmolysis only occurs in cells that are completely immersed in solution and have no air spaces around them, as in epidermal strips floating on water. Plasmolysis starts when the osmotic pressure of the solution is increased above that of the cells, causing the protoplast to shrink, and the plasma membrane separates from the wall (Fig. 3.7). Large gaps created between the plasma membrane and the wall fill with the bathing solution. This cannot occur in normal tissues as the cells have air spaces between them that are not filled with water. This includes root cells of intact plants growing in hydroponic solution or in waterlogged soil, as they still have air spaces.
Air cannot enter the cell through the cell walls as the small pore size, about 15 nm, would need a suction of 20 MPa to drain the pores (Table 3.1, in previous section) which is impossible.
During plasmolysis, the plasma membrane is stretched into strands that remain tethered to the wall at particular sites (Figure 3.8). Plasmolysis has been used by microscopists to demonstrate the tethering of the plasma membrane to specific sites on cell walls, by floating tissue such as epidermal peels of onion bulbs on high concentration of solution of sucrose (Figure 3.8). When the protoplast shrinks away from the cell wall, and solution with small molecular weight solutes pentrates the cell wall and floods into the space between the wall and the proplasts, some parts of the plasma membrane stay tethered and the rest become pulled into very fine strands.
The difference between cytorrhysis versus plasmolysis is most easily seen in leaves with single cell layers like mosses. Figure 3.9 shows that in the hydrated leaflet of Physcomytrella, when the central vacuole is distended, the chloroplasts line the cell wall. Rapid water loss causes a general shrinkage and eventually a collapse at the central parts of the cells. In cytorrhysis, the plasma membrane always remains in close contact with the cell wall. In contrast, when cells are bathed in a solution of small molecules like sucrose, glycerol, or low molecular weight polyethylene glycol, PEG, the solutes pass through the cell wall but not the plasma membrane, causing shrinkage of the protoplast and detachment of the plasma membrane from the cell wall. In plasmolysis, the gaps between the cell wall and the plasma membrane are filled with plasmolytic solution (Figure 3.9).
Cytorrhysis also occurs during freezing, when water is withdrawn from cells (Buchner and Neuner 2010).
Plasmolysis is a laboratory phenomenon and does not occur in nature. It is an experimental artifact.