2/13/2020

The University of the South Pacific

BI 302 Plant Physiology

Plant Water Relations

Key elements
• Major processes of water movement in plant cells.

• What is water potential?

• Water movement in plant cells.

• What physical forces influence water movement at

cellular level in plants?

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Uses of water in plants

• 80-95% of fresh weight of most plants.

• Solvent in which gases, minerals and other solutes dissolve

and move from cell to cell and organ to organ.

• Reactant or substrate in many reactions, including

photosynthesis and hydrolytic processes such as the amylase-
mediated hydrolysis of starch to sugar.

• Maintains turgidity, essential for cell enlargement and growth.

Water availability limits plant growth and

productivity of natural ecosystems

From: Tiaz L and Zeiger E. 2002. Plant Physiology. Sinauer Associates, Inc. Pg.34.

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Water movement in
plants is through
Translocation -
Movement of
substances from one
region to another

Water uptake and transport are governed by

physical laws
The flow of water up a
plant, driven by
Like Lego, water is very
evaporation, can be
cohesive, meaning that
replicated in a purely
the molecules stick
physical model
together and a column
of water can be pulled
“We may compare this to a great height
action to the action of a
porous vessel drawing
up a column of liquid to
supply the evaporation
loss at its surface”
– Dixon and Joly (1895)

Dixon, H.H. and Joly, J. (1895). On the Ascent of Sap. Phil. Trans. Roy. Soc. London. (B.). 186: 563-576.

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Structure and Properties of water

• Polar molecule bound together by hydrogen bonds
results into unusual properties of water.

Water has many unusual and important properties

δ- - The arrangement of atoms in a water Water’s high dielectric

molecule means that charge is constant means it is a
O distributed asymmetrically; it is polar good solvent for polar
H H + and it has a high dielectric constant compounds
δ+ δ+ Boiling point
100 Melting point
50
Water forms 0
intermolecular -50
hydrogen bonds and
so is very cohesive. It
δ- -150
sticks together and
δ+ has unusually high
-250
boiling and melting CH4 NH3 H2 O HF
δ-
δ+ points
δ- δ+
δ+ δ-
Exobiologists usually
consider water a
prerequisite for life
Photo credit: NASA; See Nobel, P. (2009) Physicochemical and Environmental Plant Physiology. Academic Press.

© 2014 American Society of Plant Biologists

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Active and Passive Transport?

Water movement in plants

• Translocation of water is a passive process.
• Diffusion and bulk flow.

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Water transport process

Diffusion
• Movement of substances due to random thermal agitation
from regions of high free energy (high concentration) to
regions of low free energy (low concentrations.

• Directed by a concentration gradient.

Fick’s law describes the movement of water by

diffusion

dm dc
DA
dt dx [] []
Fick’s Law
dc
dm/dt = amount of substance dx
moved per unit time
D = diffusion coefficient
(property of substance and
matrix) x
dc/dx = gradient of
concentration (change in
concentration per unit A = area through
of distance, dx, in a direction which flow occurs
perpendicular to the plane A)

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Diffusion rate is affected by area,

distance, and gradient
dm dc
DA
dt dx [] []

dc
Diffusion is faster when:
• The concentration dx
gradient dc/dx is steeper
• The area A is larger
• The distance x is smaller x
• D is larger

A = area through
which flow occurs

Diffusion rate is affected by the properties of

material and solvent

dm dc
DA
dt dx
The diffusion coefficient
D is affected by:
• Molecular shape
• Molecular size
• Molecular charge
• Solvent viscosity
• Solution properties as
salt concentration, pH,
temperature etc.

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Diffusion is rapid over short distances but

extremely slow over long distances
• As substances diffuse away from the starting point,
concentration gradient becomes less steep and net
movement becomes slower

Potassium permanganate crystals begin to diffuse in the water to give layers of different concentrations.

From: http://www.bbc.co.uk/bitesize/intermediate2/biology/living_cells/diffusion_and_osmosis/revision/2/

Tequila Sunrise
Pour the tequila and orange juice into glass over ice.
Add the grenadine, which will sink to the bottom.
Do not stir.
Garnish and serve.

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Osmosis
• Diffusion of water across the selectively permeable
membrane from an area of low solute concentration to one
of high concentration.
• Both concentration gradient and pressure gradients drive
transport in osmosis.
• Direction and rate of water flow across the membrane is
determined by the sum of these two driving force known as
water potential.

Water movement by diffusion of individual water molecules through the

cell membrane and by bulk flow through water selective pore (integral
protein aquaporin)

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Water potential
Water potential is a measure of free energy status
of water.

The chemical potential is the free energy per mole

of any substance in a chemical system (J mol-1).

In plant water relations the chemical potential is

referred to as water potential – free energy of
water per unit volume (J m-3).

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• Plants can use the potential energy in water to perform work.

• Water is pulled to the top of the tree because of differences in the

water potential, which is a function of the amount of dissolved
nutrients.

• Movement of water from regions of high water potential to

regions of low water potential – just another way to describe
osmosis.

Tomato plant regains turgor pressure –

cell pushes against wall due to uptake of
water

• Water potential is symbolized by w .

• Water potential of pure water is taken as zero.
• Presence of any solutes etc. will lower the energy of the
system below zero (i.e. negative).
• Plant cell growth, photosynthesis and crop productivity are
all strongly influenced by water potential and its
components.

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• There are two components to water potential:

- physical forces or pressure, such as plant cell wall or

gravity.

- the concentration of solute in each solution.

• The combined effects of solute concentration and

physical pressure (cell wall) can be measured as Water
Potential 

•  is measured in megapascals (MPa)

1 Mpa = 10 atmospheres of pressure

Osmotic forces drive the movement

of water into and out of cells
Cells have a lower osmotic Fresh water Fresh water
potential than pure water,
(because of the salts and
proteins in them), so water
moves into them An animal cell might burst Plant cell walls prevent them
from bursting

Salt water Salt water Salt water has a lower

osmotic potential than
cells, so water flows
outwards

Osmotic potential is written as Ψπ and measured in MegaPascals (MPa)

For seawater, Ψπ is about -2.5 MPa, and for a typical cell, Ψπ is about -0.8 MPa

© 2014 American Society of Plant Biologists

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Water moves down osmotic gradients

Water moves across a

semipermeable membrane
into a compartment
containing salt

The water flows inwards

until the force of the
pressure inside balances
the osmotic driving force

The osmotic potential of pure water is 0. The addition

of solutes lowers the water potential. Water moves
towards a region with a lower water potential.

© 2014 American Society of Plant Biologists

Factors contributing to cell water potential

• Solute concentration, pressure and gravity influence the
water potential.
• Hence

w = p + s + g

Where w is the water potential

p is the pressure potential
s is the solute potential
g is the gravity potential

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Solute potential
• Represents the effect of dissolved solutes.

 S of any solution at atmospheric pressure is always

negative – why?

• Solutes reduce the free energy of water by diluting

it.

Pressure potential
• P is the hydrostatic physical pressure of a solution.
• Positive pressures increase the water potential while
negative pressures reduce it.
• Positive hydrostatic pressure within the cells is
referred to as turgor pressure.

• P can be negative  transpiration in the xylem

tissue of a plant (water tension).
• P can be positive  water in living plant cells is
under positive pressure (turgid).

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Pressure can be positive or negative

Pressure Car tire Pressure
washer Household pressure required to
15 MPa water 0.25 MPa blow up a
pressure
balloon
0.3 MPa
0.01 MPa

Vacuum cleaner Inside typical plant cell Human blood

-0.02 MPa 0.5 to 1.5 MPa pressure
(household) < 0.02 MPa
-0.1 MPa
(commercial)
Laboratory Inside xylem:
vacuum From +1 MPa to
-0.01 MPa -3 MPa or lower

* These numbers are relative to atmospheric pressure (0.1 MPa), not absolute Mschel; Image 14869 CDC/ Nasheka Powell

© 2014 American Society of Plant Biologists

Turgor pressure supports plant cells and tissues

Young, non-woody
tissues are supported
by turgor pressure

Water limitation lowers

turgor pressure and
tissues wilt

Clemson University - USDA Cooperative Extension Slide Series ,R.L. Croissant, Bugwood.org

© 2014 American Society of Plant Biologists

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Gravity potential
• Water moves downward unless the force of gravity is
opposed.
• Depends on height (h) of the water above the reference
state water, density of water (pw) and the acceleration
due to gravity.
• Gravitational component is usually omitted when dealing
with water at cellular level since it is negligible compared
to P and s

• Hence at cellular levels:

w = p + s

• Water can enter of leave the cell in response

to a water potential gradient.

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Water potential of the

cell is less than the
water potential of the
solution, cell takes up
water.

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Increasing the solute concentration lowers the water potential of

the solution, hence water from the cell is drawn out, cell turgor
pressure decreases, protoplast pulls away from the cell wall.

Applying pressure removes water out of the cell hence increases the
solute potential of the cell.

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Water moves by diffusion and bulk

flow
Diffusion: Random movement of
individual molecules (1, 3)
ψw =
-100 MPa 3. Water
moves from
leaf to air

2. Water moves
through plant

Bulk flow: water in streams, tubes,

hoses, water column of xylem (1, 2)
ψw =
-0.3 MPa 1. Water
moves from
soil to root

© 2014 American Society of Plant Biologists

Diffusion rate is affected by the properties of material

and solvent

Cell membranes and walls

At cellular levels increase the diffusion
coefficient and can drastically
lower the rate of diffusion

Channels between cells called

plasmodesmata and water
channels called aquaporins can
accelerate intercellular diffusion

A small molecule can

diffuse across a 50 μm cell
in 0.6 s, but it would take 8
years to diffuse 1 m

© 2014 American Society of Plant Biologists

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Bulk flow (mass flow)

• Concerted movement of groups of molecules (along with
everything in it) in response to a external force (pressure
gradient).
• Independent of solute concentration gradients.
• Mechanism for long distance transport of water in the
xylem.

Long-distance water movement occurs by bulk flow

Bulk flow is much faster Waterfall:

than diffusion, but requires Energy
the input of energy provided by
gravity

Human
circulatory
system: Water
Energy movement
provided by up a plant:
mechanical Solar energy
pump drives
(heart) evaporation

Plastic Boy; Poco a poco

© 2014 American Society of Plant Biologists

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Water potential is an indicator of plant health

What do various values of ΨW mean for plant function?

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