Plants transport water to great heights using water potential.
Water potential is a measure of the potential energy in water.
It is the difference between the potential energy of a water sample and the potential energy of pure water.
Expressed as the Greek letter Ψ (psi).
Measured in units of pressure - megapascals (MPa).
The potential energy of pure water is set to zero, so other potentials are relative to it.
Factors Influencing Water Potential
Water potential in a plant is influenced by:
Solute concentration
Pressure
Gravity
Matrix effects: ion effects within the plant
Water Movement and Equilibrium
As each component changes, the water potential of the system raises and lowers.
Water moves to equilibrate the system, flowing down a potential gradient from an area of higher potential to lower potential.
For water to move from the soil into the plant and up to the atmosphere (transpiration):
Ψ{soil} > Ψ{root} > Ψ{stem} > Ψ{leaf} > Ψ_{atmosphere}
Water movement occurs exclusively in response to changes in water potential.
Solute Potential (Ψ_s)
Related to solute concentration, described by the van 't Hoff equation:
Ψ_s = -MiRT
M = molar concentration of the solute
i = van 't Hoff factor (ratio of particles in solution to formula units dissolved)
R = ideal gas constant
T = temperature in Kelvin degrees
Characteristics of Solute Potential
Solute potential is negative in plant cells and zero in distilled water.
Solutes reduce water potential, resulting in a negative water potential.
This is due to the bonds between solutes and hydrogen molecules; energy in hydrogen cannot do work when bonded.
Solute potential decreases with increasing solute concentration, causing a decrease in the system's water potential.
Pressure Potential (Ψ_p)
Pressure is an expression of energy.
The higher the pressure, the more potential energy in a system.
Positive pressure potential (compression) increases the system's water potential.
Negative pressure potential (tension) decreases the system's water potential.
Plant Manipulation of Pressure Potential
Plants manipulate pressure potential by adjusting solute potential and osmosis.
↑solute concentration = ↓solute potential = ↓total water potential = ↑water into cell = ↑pressure potential
The opening and closing of stomata also influences pressure potential by releasing water from the plant.
Gravity Potential (Ψ_g)
In a plant with no height, gravity potential is close to negative 0.
Gravity always reduces the water potential of the system by pulling downward towards the soil.
The taller the plant, the higher the water column, and the greater the influence of gravity potential.
Plants cannot manipulate gravity potential.
Matric Potential (Ψ_m)
Always negative - 0, and 0 in a water-saturated system.
The binding of water to a matrix always removes potential energy from the system.
Similar to solute potential in binding water to another component, such as insoluble, hydrophilic molecules of the cell wall.
Very (negatively) large in dry seeds; decreases as seeds hydrate.
Cannot be manipulated by the plant.
Typically ignored in well-watered plant systems because it is close to zero.
Transpiration
The loss of water through evaporation at the leaf surface.
Main driver of water movement through xylem.
Causes negative pressure (tension) at the leaf surface, varying due to atmospheric humidity.
During transpiration, water is drawn from the leaf mesophyll cells, through the xylem, down the stem, through the root cells and root hairs, and into the soil.
Stomata close at night, and water is stored in the xylem vessels and tracheids by cohesion and adhesion.
Passive process - NO ATP is required.
Control of Transpiration
Atmosphere drives transpiration rate and causes massive water loss (up to 90% of water taken up).
Cuticle: a waxy covering on leaves that prevents some water loss.
Stomata & guard cells open and close to regulate gas exchange & water loss.
Adaptations to Water Loss
Thick waxy cuticle to prevent water loss, as seen in desert plants or xerophytes.
Thick covering of trichomes or stomata under the leaf surface reduces water loss during transpiration.
Transportation of Photosynthates in the Phloem
Photosynthates: the products of photosynthesis, typically in the form of simple sugars like sucrose.
Sources: structures in the plant that produce photosynthates (leaves).
Translocation: transporting photosynthates to the growing parts of the plant.
Sinks: points for sugar delivery; e.g., roots, shoots, developing seeds.
Photosynthates are sent to different parts of the plant during different stages of development.
Sugars from leaves may be sent out to the roots for storage, or sugars in storage in the roots may be sent to leaves.
Translocation Process
Photosynthates are produced in the mesophyll of leaves, connected by plasmodesmata.
Sugars are transported to reach phloem sieve tube elements (STE), AGAINST its concentration gradient (ATP required).
STE’s have reduced cytoplasmic material & are separated by porous plates, which allow for pressure-driven bulk flow.
Movement to Closest Sink
Photosynthates are moved to the closest sink.
Phloem sap is an aqueous solution containing ~30% sugar, minerals, amino acids, and plant growth regulators.
High sugar content decreases solute water potential, thereby decreasing total water potential, which drives water into the phloem from the xylem, increasing pressure.
Increased pressure drives bulk flow because sink concentration is lower than the phloem.