Plants, Cells, and Water

Chapter 1
 Learning objectives
Know the unique properties of water and
their biological importance.

Understand the meaning of the terms

adhesion, cohesion, and tension.

Understand the processes of diffusion and

bulk flow as mechanisms of water
transport.
 Learning objectives
Understand the process of osmosis as a
process of water movement across a
selectively permeable membrane.

Understand the concept of water potential

and its component potentials.

Know the role of aquaporins in the

transport of water.
 Plants, cells, and water
The properties of water allow this molecule to
fulfill specific roles in plants.
The thermal properties of water provide temperature
regulation and maintain water in a liquid state.
Water serves as the universal solvent.
The hydrolysis or condensation of water can
contribute to biochemical reactions.
The transparency of water does not hinder
photosynthesis.
Water creates turgor pressure.
Physical and chemical properties of water

The electronegativity of the oxygen atom in

water makes the molecule polar.

The partial negative charge on the oxygen atom

and the partial positive charge on the hydrogen
atoms allows water to form hydrogen bonds.

This polarity also allows water to interact with

other molecules and surfaces, creating layers of
water molecules called bound water.
Physical and chemical properties of water

Figure 1.1
Physical and chemical properties of water

The thermal properties of water are key to

its role in biological systems.
Water has a high specific heat, which means
it has a high capacity to absorb heat.
Water has high thermal conductivity, which
allows it to conduct heat away from the site
where the heat is generated.
The heat of fusion for the conversion of ice
to water is high.
Physical and chemical properties of water

Table 1.1
Physical and chemical properties of water

The thermal properties of water are key to

its role in biological systems.
The density of water is highest at 4 C rather
than at the freezing point (0 C).
The heat of vaporization of water, which is
the energy required to covert liquid water to
water vapor, is also high.
The high dielectric constant contributes to
waters ability to form hydration shells
around molecules.
Physical and chemical properties of water

Figure 1.2
Physical and chemical properties of water

Table 1.2
Physical and chemical properties of water

The polarity of water molecules creates a

strong attraction between water molecules
called cohesion.

The cohesive properties of water directly

contribute to waters tensile strength and
surface tension.

The polarity of water also creates a strong

attraction to surfaces called adhesion.
Physical and chemical properties of water

Figure 1.3
 The movement of water
The transport of water occurs by the
energetically passive processes of
diffusion and bulk flow.
Bulk flow (or mass flow) is driven by
differences in pressure.
Diffusion is driven by difference in the
concentration of water.
 The movement of water
Diffusion is governed by Ficks first law.
1
J = -D A
C l
J is the flux (amount of material crossing per unit area
per unit time.
D is the diffusion coefficient of the media.
A is the cross-sectional area.
L is the length of the diffusion path.
C represents the concentration gradient.
The negative sign indicates that diffusion only occurs
from a higher concentration to a lower concentration.
 The movement of water
Figure 1.4
 The movement of water
Osmosis is a special case of diffusion
involving the movement of water across a
selectively-permeable membrane.

A selectively permeable membrane allows

for the movement of water, but restricts the
movement of other solutes.
 The movement of water
Figure 1.5
Selectively permeable membranes
in plant cells
Plant cells have a variety of selectively
permeable membranes that create various
subcellular compartments.
The outer plasma membrane encloses the
cells liquid protoplasm.
The internal cellular structures, called the
organelles, are either composed of, or
surrounded by, membranes.
In addition to the various membranes, the cell
is encased with a cell wall.
Selectively permeable membranes
in plant cells
Figure 1.6
Thermodynamics of water movement

The passive movement of water or other

substances across a membrane is
dictated by the chemical potential () of
that substance on each side of the
membrane.
The chemical potential is the free energy per
mole of a substance.
The chemical potential also is a measure of
the capacity of a substance to move.
Thermodynamics of water movement
Pure water has a chemical potential of zero.

The addition of solutes decreases the mole

fraction of water, calculated by:
w
XW
w+s
Xw is the mole fraction of water
W is the moles of water
S is the moles of solute
Thermodynamics of water movement

Using the mole fraction of water, the

chemical potential for water on each side
of the membrane can be calculated by:
w RT ln X w
w
*

w is the chemical potential of water on a

given side of the membrane
w* is the chemical potential of water under
standard temperature and pressure
Xw is the mole fraction of water
Thermodynamics of water movement

For two sides of a membrane with different

chemical potentials, water will move
passively from the higher chemical
potential to the lower chemical potential.

Since water only moves passively, plants

control the movement of water indirectly
by moving ions to alter the w.
Thermodynamics of water movement

The movement of ions to control the

movement of water may require that the
ions be moved against their gradient,
which is an example of active transport.

The manipulation of ion transport to

control water transport is an example of
osmotic adjustment.
Thermodynamics of water movement

Figure 1.7
Thermodynamics of water movement
The effect of ions on the osmotic component of
the chemical potential of water can be
demonstrated with an osmometer.

When water moves down its chemical potential

gradient across a membrane, it will continue to
do so until an equilibrium is reached.

If a piston applies positive pressure to exactly

balance (and prevent) the flow of water, this
pressure is called the osmotic pressure
Thermodynamics of water movement

The magnitude of osmotic pressure

needed to prevent the osmosis of water is
directly related to the solute concentration.

Since the osmotic pressure offsets the

potential for water movement, the solute
potential (S) of the solution is the
opposite sign of the osmotic pressure.
W -
Thermodynamics of water movement

Figure 1.8
Thermodynamics of water movement

As the osmometer illustrates, the chemical

potential also has a pressure potential
(VP) such that:
w RT ln X w+ VWP
w
*

There are also electrical and gravitational

components to the chemical potential, but
for water these are generally negligible, so
they are omitted from the equation.
Thermodynamics of water movement

By substituting values in the equation, and

including both the osmotic and hydrostatic
pressure, the chemical potential of water
can be described as:
w VW(P- )
w
*

And with some final substitutions the

water potential of a solution is illustrated
as: P-
Thermodynamics of water movement

Water potential is a useful measurement

for plant physiologists because:
P and can be measured experimentally.
The units for water potential (pascals) are
relevant to the soil-plant-atmosphere system.
Gradients in water potential also describe the
passive flow of water, specifically from more
positive to more negative values.
Water potential can be used as a measure of
physiological status.
Thermodynamics of water movement
The water potential can also be expressed as
the sum of the component potentials.

P+ S
P is the pressure potential (same as P)
S is the solute (or osmotic) potential

The matric potential is sometimes also included,

which accounts for the adsorption of water to
solid surfaces, such as in seed imbibition.
Thermodynamics of water movement

In plant cells, the vacuole makes the

greatest contribution to the water potential
because the vaciole contains dissolved
solutes and 50-80% of the cellular water.

When plant cells gain water, they exert an

outward pressure called turgor pressure.

A plant cell full with water is turgid.

Thermodynamics of water movement

When turgid plant cells exert turgor

pressure against the wall, there is an
equal, opposite wall pressure.

If the cell loses water to the point that the

turgor pressure is zero, the cell is flaccid.

Incipient plasmolysis refers to the

condition where the protoplast is filled with
water and turgor pressure is zero.
Thermodynamics of water movement

In a hypotonic solution the cell would

gain water because cell is more negative
than the solution.

In a hypertonic solution the cell would

lose water and undergo plasmolysis
because cell is more positive than the
solution.
Thermodynamics of water movement

Water potential gradients also determine

the flow of water from one cell to another.

In nature, an excessive loss of water from

plant cells leads to wilting.

Under conditions of wilting, the water

potential of the cell becomes highly
negative.
Thermodynamics of water movement

Figure 1.9
 Aquaporins
Porins are major intrinsic proteins
(MIPS), a class of membrane proteins.

Porins are non-selective cation channels.

Aquaporins are found in the plasma

membrane and tonoplast and mediate the
transport of water and neutral solutes like
glycerol.
 Aquaporins
Figure 1.10a
 Aquaporins
The structure of aquaporins is very similar
across different organisms.

Aquaporins are formed from four protein

subunits.

Aquaporins are gated channels, which

means that they can be selectively opened
and closed.
 Aquaporins
Figure 1.10b
 Aquaporins
Aquaporins are not the only mean by
which water can enter cells.

Water can diffuse directly through the cell

membrane between the phospholipid
molecules.

Aquaporins provide a more rapid rate of

water movement, and therefore make a
greater contribution to osmoregulation.
 Aquaporins
Figure 1.10c
 Osmoregulation
Osmoregulation allows cells to respond to
the external environment.

An osmoregulatory system involves a two-

component system:
A sensing mechanism (sensor protein or
osmosensor)
A transducing mechanism (response
regulators such as transcription factors)
 Osmoregulation
Figure 1.11
 Osmoregulation
In E. coli, the two-component system
consists of:
The EnvZ sensor protein
The OmpR response protein

The system regulates the expression of

the pore proteins OmpF and OmpC.
 Osmoregulation
In yeast, the two-component system has
SLN1, a histidine kinase, as the
osmosensor.

A similar system is present in Arabidopsis

thaliana with ATHK1 as the osmosensor.