Plant Water

Relations and Water

Economy
Chapter 6
General Cell Characteristics
• Plant cell ~ 90% water
• Vacuole ~ 80-90% of cell volume
• Most free water is in vacuole
• Vacuole water is used to buffer and
regulate cytoplasmic moisture content
• Cytoplasmic moisture must be regulated
within strict limits
 The Energy in Water
• All molecules have energy and are in
motion
Without stirring
 A Law of Nature?
• All molecules and substances tend to
reach their lowest energy level
– Rocks roll downhill
– Water seeks the lowest elevation
possible
– Air molecules will slow unless heated

Therefore, water will move to an area

where its energy level is lower.
 Osmosis
The movement of water through a differentially
permeable membrane, where water can cross but salt
cannot

Osmotic Potential
Forces causing water to move across a
differentially permeable membrane
How is the energy content of water
defined?

Water’s chemical potential ( µ ) is the

“free energy” per mole.
What factors govern chemical
potential? Concentration (actually
activity), temperature, pressure,
electrical potential (zero for water),
and gravity.

Although chemical potential defines

water’s energy, plant physiologists use
a concept called water potential (  ).
 How is the concentration of solute
determined? Not directly by mass.
1 Gram Molecular Weight (GMW) =
Atomic weight of a molecule converted to
grams
1 GMW/1000g of H2O = 1.0 molal solution
1.0 molal solution = -22.7 bars of pressure
0.5 molal solution = -11.35 bars
1 bar = 1 atmosphere of pressure
 Sugar Example
• Sucrose = C12H22O5
• Atomic Weights: Therefore, 342 g L-1 is 1 M.
C = 12 However, if the cell inverts
sucrose to glucose and fructose
H=1 it acts like 2 M.
O = 16
• 1 GMW of sucrose = 342 g
 Salt Example
• Salt = NaCl
Therefore, 58.5 g L-1 is 1 M.
• Atomic Weights: However, NaCl ionizes so
solution acts more like 2 M.
Na = 23
Cl = 35.5
• 1 GMW of sodium chloride = 58.5 g
 Temperature Effects

• Temperature directly affects that energy

and the degree of motion
• A solute (dissolved substance) attracts
water molecules and:
– slows the motion of water molecules
– reduces the energy level of water
molecules
 = Water Potential
 = Osmotic Potential
p = Turgor Potential

• Pure water has the highest energy and is

defined as  = 0.00
• Add sugar or salt and  becomes
negative
 Expression relating water potential () to
osmotic and turgor variables

 = hydrostatic pressure - osmotic pressure

 = pressure potential + osmotic potential

 = p + 

osmotic potential is sometimes called solute potential

 Turgor Pressure
• A result of the vacuole being full of water
and pressing the cytoplasm (and plasma
membrane) against the cell wall (turgid).
• A cell becomes flaccid when the vacuole
is forced to give up too much water to the
cytoplasm. This results in leaf and plant
wilt.
• Vacuole acts as the buffer, providing
water upon demand.
 How is cytoplasmic water
regulated?

• Vacuole acting as a buffer reservoir

• Guard cells open/close depending on
water relations and other factors (see
earlier notes on photosynthesis and
growth and notes later in this lesson)
 Water Movement
• Water will always move from an area of

High Low Water potential

0.00  negative 
Less negative more negative 
High energy low energy
What determines whether water from one
side of a membrane will cross that
membrane?
Permeability Equation, Fick’s first law
J = - D A C / l

Besides the membrane permeability factor, the

difference in the concentration of water otherwise
called the mole fraction of water is the key factor.
Pure water is 55.5 molal. Water will move from the
side of greater chemical potential to lower chemical
potential.
 Summary of Water
Movement Forces
• Water enters vacuole of cell
• Vacuole presses cytoplasm against cell
wall
• Turgor pressure builds
• Water uptake by vacuole stops when cell
wall pressure = 
• Vacuole regulates  of cytoplasm
There is an additional “force” that is
often include in discussions of plant
water relations. The matric potential
relates to the affinity of the cell wall
or other structural component for the
water. A seed is an example where
matric potential would be strong and
would reduce water potential.
 What Happens Under
Water Stress?
• Water evaporates through stomates
• Under stress, it is difficult to replace
from soil through the root system
• Xylem and cell wall space under
tremendous vacuum (-)
• Water leaves cytoplasm of leaf cells
• Cytoplasm water replaced from vacuole
• Plant wilts
• When vacuole is depleted, reach PWP
flaccid

p= cell -

p= 0, = cell
Other methods used to determine water potential

1. Scholander Pressure Bomb (Fig. 6-16)

2. Psychrometers
(water and osmotic potential)

3. Relative Water Content (not really water

potential
RWC = (fresh - dry) / (turgid- dry)
 Winter Injury
• Water in cell wall space begins to freeze
• Solutes are concentrated in remaining
liquid water (which reduces )
• Water leaves cytoplasm to cell wall space
• Vacuole is depleted
• Cytoplasm becomes too concentrated
• Cell dies
 Water Absorption From Soil
• Water absorbed through epidermal cells
near root tip (mostly through root hairs)
• By osmosis
• Water movement from soil (low ) to
epidermal cell (lower )
•  ~ 0.0 in a soil at field capacity
• As soil dries,  decreases (more negative)
• When soil  becomes more negative than
cell , plant wilts
Soil Type vs. Water Relations
% Water
Soil Type FC Wilting % % Avail.
H2O
Fine Sand 3.3 1.3 2.0
Sandy Loam 18.5 10.0 8.5
Silt Loam 21.3 10.4 10.9
Clay 28.0 14.5 13.5
 Water Movement
Through a Plant
Root Hair Cortex Through Cells
“Symplast”

To Xylem *Between Cells*

non-living, “Apoplast”
empty cells
Endodermis or
Through cells Casparian Strip
Typical Example of  Values
(in bars)

Soil -0.5
Root -2
Stem -5
Leaf -15
Air -1000
 Transpiration
• Stomates: There are between 8,000 and
60,000/cm2
• Leaves exposed to the sun have most
stomates
• 1-3% of leaf surface is stomates
Resistances to Gas Exchange

• Mesophyll and Substomatal Cavity

• Stomatal
• Boundary Layer
Transpiration Rate Depends on:
• Size of stomatal opening
• Difference in  of leaf and atmosphere
(correlated to vapor pressure deficit). VPD is
affected by temp and humidity.
• Wind - reduces boundary layer resistance

Examples:
• 100% RH = 0.00 
• 50% RH = several hundred bars
• Leaf is seldom less than -20 bars
 Water Vapor Pressure
Relative Humidity
Temp °C 100% 80% 50% 20% 10%
----------------------- kPa -----------------------
30 4.24 3.40 2.12 0.85 0.42

20 2.34 1.87 1.17 0.47 0.23

10 1.23 0.98 0.61 0.24 0.12

Guard Cell Control
Opening Vacuole
Process

H2O

H
+ K+
Closing
Process H2O

K+
H+
 Stomatal Operation
• Once thought that wilting closed stomates but
now known that abscisic acid (ABA) increases,
causing a decrease in K+, a loss of water, and
stomate closing
• Other controls
– low CO2 opens
– high CO2 closes
– high temp closes
– dry soil closes
– low humidity opens
 Note:
• A biological clock mechanism also
controls stomatal opening/closing

• Example: if plants are artificially kept in

the dark from the previous night,
stomates will open at the time of normal
sunrise
 H-Bonding
• Xylem water is normally under
tremendous tension (vacuum)
• H-bonding is critical to maintaining the
continuity of the column of water in
xylem tissues
• Xylem tissues often have internal spiral
wall thickening to provide strength
Hydrogen Bonding
 Root Pressure
• Some mineral elements are actively
transported from root into xylem
• If transpiration rate is low, then salts
accumulate in xylem and  is reduced
• Water is osmotically moved to the xylem
• Pressure builds, and xylem contents have to
go somewhere
• Results in:
– guttation
– excessive stem bleeding (ex. milkweed)
Guttation
 Why do plants transpire when such
behavior prevents them from
growing under dry conditions?

Possible Reasons:
1. Transpiration flow is needed for mineral transport.
However, plants transpire very dilute aqueous solutions
during high temperature conditions.
2. Transpiration helps to cool the leaves. The heat of
vaporization of water is extremely high and helps to
prevent leaves from overheating.