第30章植物の形態と生理

Pressure potential (Ψ_p), also called turgor potential, may be positive or negative (Figure 30.32). Because pressure is an expression of energy, the higher the pressure, the more potential energy in a system, and vice versa. Therefore, a positive Ψp (compression) increases Ψ_total, and a negative Ψ_p (tension) decreases Ψ_total. Positive pressure inside cells is contained by the cell wall, producing turgor pressure. Pressure potentials are typically around 0.6–0.8 MPa, but can reach as high as 1.5 MPa in a well-watered plant. A Ψ_p of 1.5 MPa equates to 210 pounds per square inch (1.5 MPa x 140 lb/in^-2 MPa^-1 = 210 lb/in^-2). As a comparison, most automobile tires are kept at a pressure of 30–34 psi. An example of the effect of turgor pressure is the wilting of leaves and their restoration after the plant has been watered (Figure 30.33). Water is lost from the leaves via transpiration (approaching Ψ_p = 0 MPa at the wilting point) and restored by uptake via the roots.

A plant can manipulate Ψ_p via its ability to manipulate Ψ_s and by the process of osmosis. If a plant cell increases the cytoplasmic solute concentration, Ψ_s will decline, Ψ_total will decline, the ΔΨ between the cell and the surrounding tissue will decline, water will move into the cell by osmosis, and Ψ_p will increase. Ψ_p is also under indirect plant control via the opening and closing of stomata. Stomatal openings allow water to evaporate from the leaf, reducing Ψ_p and Ψ_total of the leaf and increasing Ψ between the water in the leaf and the petiole, thereby allowing water to flow from the petiole into the leaf.

Gravity potential (Ψ_g) is always negative to zero in a plant with no height. It always removes or consumes potential energy from the system. The force of gravity pulls water downwards to the soil, reducing the total amount of potential energy in the water in the plant (Ψ_total). The taller the plant, the taller the water column, and the more influential Ψ_g becomes. On a cellular scale and in short plants, this effect is negligible and easily ignored. However, over the height of a tall tree like a giant coastal redwood, the gravitational pull of –0.1 MPa m^-1 is equivalent to an extra 10 MPa of resistance that must be overcome for water to reach the leaves of the tallest trees. Plants are unable to manipulate Ψ_g.

Matric potential (Ψ_m) is always negative to zero. In a dry system, it can be as low as –2 MPa in a dry seed, and it is zero in a water-saturated system. The binding of water to a matrix always removes or consumes potential energy from the system. Ψ_m is similar to solute potential because it involves tying up the energy in an aqueous system by forming hydrogen bonds between the water and some other component. However, in solute potential, the other components are soluble, hydrophilic solute molecules, whereas in Ψ_m, the other components are insoluble, hydrophilic molecules of the plant cell wall. Every plant cell has a cellulosic cell wall and the cellulose in the cell walls is hydrophilic, producing a matrix for adhesion of water: hence the name matric potential. Ψ_m is very large (negative) in dry tissues such as seeds or drought-affected soils. However, it quickly goes to zero as the seed takes up water or the soil hydrates. Ψ_m cannot be manipulated by the plant and is typically ignored in well-watered roots, stems, and leaves.

Solutes, pressure, gravity, and matric potential are all important for the transport of water in plants. Water moves from an area of higher total water potential (higher Gibbs free energy) to an area of lower total water potential. Gibbs free energy is the energy associated with a chemical reaction that can be used to do work. This is expressed as ΔΨ.

Transpiration is the loss of water from the plant through evaporation at the leaf surface. It is the main driver of water movement in the xylem. Transpiration is caused by the evaporation of water at the leaf–atmosphere interface; it creates negative pressure (tension) equivalent to –2 MPa at the leaf surface. This value varies greatly depending on the vapor pressure deficit, which can be negligible at high relative humidity (RH) and substantial at low RH. Water from the roots is pulled up by this tension. At night, when stomata shut and transpiration stops, the water is held in the stem and leaf by the adhesion of water to the cell walls of the xylem vessels and tracheids, and the cohesion of water molecules to each other. This is called the cohesion–tension theory of sap ascent.

Inside the leaf at the cellular level, water on the surface of mesophyll cells saturates the cellulose microfibrils of the primary cell wall. The leaf contains many large intercellular air spaces for the exchange of oxygen for carbon dioxide, which is required for photosynthesis. The wet cell wall is exposed to this leaf internal air space, and the water on the surface of the cells evaporates into the air spaces, decreasing the thin film on the surface of the mesophyll cells. This decrease creates a greater tension on the water in the mesophyll cells (Figure 30.34), thereby increasing the pull on the water in the xylem vessels. The xylem vessels and tracheids are structurally adapted to cope with large changes in pressure. Rings in the vessels maintain their tubular shape, much like the rings on a vacuum cleaner hose keep the hose open while it is under pressure. Small perforations between vessel elements reduce the number and size of gas bubbles that can form via a process called cavitation. The formation of gas bubbles in xylem interrupts the continuous stream of water from the base to the top of the plant, causing a break termed an embolism in the flow of xylem sap. The taller the tree, the greater the tension forces needed to pull water, and the more cavitation events. In larger trees, the resulting embolisms can plug xylem vessels, making them nonfunctional.

Transpiration—the loss of water vapor to the atmosphere through stomata—is a passive process, meaning that metabolic energy in the form of ATP is not required for water movement. The energy driving transpiration is the difference in energy between the water in the soil and the water in the atmosphere. However, transpiration is tightly controlled.