Similarly, water moves within a plant, or from wet soil into the roots of a plant, because

the movement is energetically downhill. In the place of gravity in the ball-and-hill example is a
gradient of water potential that must exist between two adjacent cells or between soil and
roots to cause water to move. It is a difference in water potential that causes water to move
from a region of high water potential (e.g. wet soil) to a region of lower water potential (e.g.
root xylem). Xylem is the network of pipe-like cells that connect roots to leaves via the stem.
The movement of water through the xylem of a plant is termed sapflow. The concept of water
potential is based on the concept of free energy. Free energy is defined as the energy available
to do work (e.g. lifting water up a tree constitutes work). Work is done when something moves
against gravity or against some other force that opposes that movement. The chemical
potential of water is defined as the free energy per mole of water and water potential is the
chemical potential of water divided by the partial molar volume of water (i.e. the volume
occupied by 1 mole of liquid water at standard temperature and pressure). Thus water
potential is a measure of the energy available for doing work, expressed per mole of water. This
rather esoteric definition is less important than the need to understand two things: first, water
moves in the SPAC from regions of high (close to zero) to regions of low (more negative) water
potential; and second, that the water potential of a plant cell has two principal components,
namely solute potential and turgor potential. Water potential can be zero or negative and the
reason for this is explained below. The meanings of solute and turgor potential are discussed
shortly.

Water potential is given the symbol "Ym (pronounced Psi W), turgor potential (a measure
of the hydroastatic or water pressure in a cell) has the symbol Y p andsolute potential (a measure
of the concentration of slotes in cell) has the symbol Ys. At this stage we can note that :

Yw = Yp + Ys

The water potential of water in a root or in a leaf, or of any system, is measured relative
to an internationally agreed reference value. The reference value has been arbitrarily defined as
the water potential of pure water at sea level, sitting in a beaker at standard temperature and
pressure (a temperature of 25°C and 0.1 MPa atmospheric pressure) and has been arbitrarily
given a water potential equal to zero. Within a plant (and for soil and atmosphere too) all other
water potentials are either lower than this value (i.e. negative) or equal to this (i.e. zero).

The reduction in free movement of a proportion of water molecules is said to reduce the ability
of the water to do work, that is, there is less energy available to do work, and therefore water
potential is reduced.
 As the concentration of solutes in the water increases, the water potential of the
solution decreases further. To a good approximation in dilute solutions, the reduction in water
potential can be calculated from:

Yw = - R X T X C

where R is the gas constant, T is the temperature (in Kelvin) and C is the osmolal concentration
of the solution. Molality is similar to molarity, but molality is a measure of the amount of
solutes present per kilogram of solvent, while molarity is a measure of the solute content per
lite of solution. Because the presence of salts (solutes) has reduced the free energy of the water
in the salty beaker, we can say that the solute potential of the water has decreased. Solute
potential (Ys) is also called osmotic potential because changes in osmotic potential of cells can
influence the osmotic movement of water between cells. When solutes (e.g. salts) are present
in water, the concentration of water molecules, and hence the free energy per volume of
water, is reduced and therefore so is water potential. Furthermore, the electrostatic interaction
between salt ions and water molecules restricts the free movement of the water molecules and
so their free energy is reduced. A definition of solute potential is therefore:

Solute potential is a measure of the change in water potential of a system due to the
presence of solutes. The more solute molecules are present, the lower (and more
negative) is Ys

Solute potential is always zero (no solutes present) or negative (solute present). The reduction
in water potential (Y,) of the salty beaker due to the presence of solutes is due to a reduction in
the solute potential (,) of the solution. Thus, water potential and solute potential are equal in
the salty beaker above. Figure 2.1 schematically outlines the situation in the two beakers. In the
beaker of salty water, water potential and solute potential are equal.

Within a plant cell, water is generally under considerable pressure. This pressure can exist
because of the presence of a strong cell wall comprised of cellulose microfibrils pushing
inwards. The air pressure in an inflated car tyre is typically in the region of 0.2-0.3 MPa (2-3 bar
or 28-42 psi air pressure). plant cell can generate hydrostatic pressures of up to 2-3 MPa. This
hydrostatic pressure is called turgor potential (y,) or pressure potential, when we are talking
about water in plant cells. Turgor potential is usually zero (e.g. in a flaccid cell in a wilted leaf) or
positive (e.g. in a turgid cell in a turgid leaf).
Thus, water potential of plant tissues has two main components - solute potential and turgor
potential:

Yw = Ys + Yp

These are the two main components of importance when considering water in plants. Later, we
shall see that an additional component, matric potential (Ym), is added in relation to soils.

The matric potential resulting from the interaction of water and surfaces (such as cell walls,
membranes and proteins) also contributes to water potential of plant cells but only becomes
significant when the water content is so low that cell death is imminent. It is usually ignored in
practical measurements. Matric potentials are sometimes considered to be accounted for in
measurements of solute potential. Matric potentials for plant tissues are rarely cited in plant
physiology or ecophysiology texts.

Water potential is zero or negative, solute potential is zero or negative and turgor potential is
zero or positive, in living cells. The unit for all potentials is Pascals (but usually in multiples of
millions, hence MPa; note that 1 MPa = 10 bar = 140 psi). When water potential of a cell is zero
the solute potential is equal, but of opposite sign, to the turgor potential of the cell. Thus in a
turgid cell the solute potential could be -1.2 MPa, the turgor potential could be 1.2 MPa and the
water potential is therefore zero (equation 3). As a cell loses water, its turgor potential declines
towards zero (the cell deflates) and its solute potential gets more negative because the solute
concentration of the cell increases. Consequently water potential of the cell declines. The
relationships among water content, water, solute and turgor potentials are discussed in depth
in Chapter 4.

OSMOSIS
Defining osmosis

Osmosis is the process by which cells exchange water with their environment. It is a passive
process similar to diffusion, but involving only water molecules. In biology, osmosis is often
defined as: the net movement of freely moving water molecules from a region of their higher
concentration to a region of their lower concentration through a partially permeable
membrane.

In the definition, a partially permeable membrane refers to a membrane that is permeable to

water and to certain solutes.

Water potential

The pressure exerted by freely moving water molecules in a system is called the water
potential, measured in kilopascals (kPa). By convention, the water potential of pure water is
given the value of O kPa. A solution with a high water potential has a high number of freely
moving water molecules.

Consider a solution with a higher water potential separated from a solution with a lower water
potential by a partially permeable membrane. More water molecules move from the region of
higher water potential to the region of lower water potential than in the other direction (figure
la). The movement continues until the water potentials on both sides of the membrane are
equal, Water molecules continue to move in both directions after equalisation, but there is no
net movement (figure 1c).

The effect of solutes

The water potential of a solution falls when solutes are added because water molecules cluster
around the solute molecules (figure 1b). As the concentration of solutes increases, clustering
also increases, further lowering the water potential.

The contribution of solutes to the water potential of a system is called the solute potential of
that system. Because it always lowers water potential, solute potential is always negative. It
becomes more negative as more solutes are added to the system