PLANT WATER RELATIONS

Net flow in
whole plants
 Some Key Concepts
• Diffusion: movement of molecules from
high to low concentration.
• Osmosis: diffusion across a semi-
permeable membrane.
• Mass or bulk flow: movement of fluid due
to pressure or gravity differences.
Water Movement
Water movement is considered to be almost entirely passive
with water flow following a water potential gradient.
Water potential is the difference in free energy of water
in soils, cells or atmosphere and that of pure water.
Water potential of pure water is used for comparison and is
given a value of zero. When there are differences in water
potential, water will ALWAYS move passively from higher
to lower water potential.
Equation for water potential (must account for the factors
that influence the diffusion of water and other
substances):
w   = p + s
where w = water potential; p = pressure potential; s =
solute or osmotic potential
1. Solute (or osmotic) potential (s).
    This is the contribution due to dissolved
solutes. Solutes always decrease the free energy of
water, thus there contribution is always negative.
2. Pressure (or Pressure Potential)(p)
    Due to the pressure build up in cells thanks to
the wall. It is usually positive, although may be
negative (tension) as in the xylem.
 WATER POTENTIAL
• Is the chemical potential of water.
• It represents free energy of water per unit volume
of water.
• Three major factors influence the water potential
in plants viz.
• Concentration,
• Pressure, and
• Gravity
 Ψ w = Ψs + Ψp + Ψ g

• Where,
• Ψw = water potential
• Ψs = solute potential
• Ψp = pressure potential
• Ψg = gravitational potential
• Solute
• Effect of dissolved solutes on water potential
• Solutes reduce free energy of water

• Pressure
• It is the hydrostatic pressure of the solution.
• Positive pressure raises the water potential while
negative pressure decreases it.
• Positive pressure – turgor and negative pressure -
tension

• Gravity
• Is generally negligible.
• Overcome by transpirational pull.
 Osmotic pressure
• Osmotic pressure
• ∏(pi)= CRT
• ∏ is the osmotic pressure
• C is molar concentration (pressure is the result of total activity of the
number of molecules)
• R is gas constant (0.082 litre atmosphers/mole degree or 0.0357 litre
cal. Per degree centrigrate)
• T is absolute temperature.
A pressure of 1 atm can also be stated as:
≡1.01325 bar
≡ 101325 pascal (Pa) or 101.325 kilopascal (kPa)
≡ 1013.25 millibars (mbar, also mb)
≡ 760 torr [B]
≈ 760.001 mm-Hg, 0 °C, subject to revision as more precise measurements of mercury’s
density become available [B, C]
≈ 29.9213 in-Hg, 0 °C, subject to revision as more precise measurements of mercury’s
density become available [C]
≈ 1.033 227 452 799 886 kgf/cm²
≈ 1.033 227 452 799 886 technical atmosphere
≈ 1033.227 452 799 886 cm–H2O, 4 °C [A]
≈ 406.782 461 732 2385 in–H2O, 4 °C [A]
≈ 14.695 948 775 5134 pounds-force per square inch (psi)
≈ 2116.216 623 673 94 pounds-force per square foot (psf)
 Problem
• Osmotic chamber contains 0.45 M sucrose
solution what will happen if it immerse in
a. pure water; b. 0.25 M solution; c. 0.45
M solution.
• Given 1 atm = 76 cm
• Specific gravity of mercury= 13.6 cm
 The Colloidal System
• Protoplasm composed of substances in its
colloidal state.
• Enzymes are active only in their colloidal
state.
• In colloidal stage 100 or 1000 of molecules
lumped together.
• Dispersed particle found in the range of 1-
100 micro meter diameter.
EXAMPLE 1: let’s suppose we
drop a plant cell into pure water

Living
Plant Cell

Water can move

by osmosis across
the cell wall and
cell membrane but
most solutes Pure water
cannot
 Plant Cell: before
equilibrating with
water
s = -0.2 MPa
p = 0 What is the total
w = ? water potential of
the plant cell?

What will happen

Pure water to the total water
potential of the
s = 0 plant cell when it
p = 0 is dropped in
water?
w = s + p = 0
 Pure water

s = 0
p = 0
w = s + p = 0
s = -0.2 MPa
p = +0.2 MPa
w = 0

This is what produces turgor, or positive pressure, in

plant cells
EXAMPLE 2: Putting a plant cell
into salty water

Plant Cell

Water can move

by osmosis across
the cell wall and
cell membrane but
most solutes salty water
cannot
 Plant Cell: before
equilibrating with
salty water
s = -0.2 MPa
p = 0
w = -0.2 MPa What will happen
to the total water
potential of the
plant cell when it
Salty water is dropped in
salty water?
s = -0.2 MPa
p = 0
w = s + p = -0.2 MPa
HKALE 1997 Biology Paper I
7. (a) Give a word equation to illustrate the relationship between the water potential (Ψ), the
osmotic potential (Ψs) and the pressure potential (Ψp) in a plant cell.(1 mark)
 
(b) Sketch a graph to show the changes inΨ, Ψs andΨp when a fully plasmolysed plant cell is
placed in pure water and regains full turgor in 40 minutes (given : pressure potential at full turgor
is +400 kPa and incipient plasmolysis occurs at 20 minutes). (5 marks)

7. (a) water potential = osmotic / solute potential + pressure potential (1)

 
(b) 1 mark for each correctly drawn curve showing the following trends, each curve must be
labelled or keyed : (total 3 marks) 3
For Ψp : rise
from 0 kPa at incipient plasmolysis at 20 minutes to +400 kPa at 40 minutes
For Ψs : rise from ＜ -
400 kPa at 0 minutes to -400 kPa at 40 minutes
For Ψ : rise from ＜ -400
kPa at 0 minutes to 0 kPa at 40 minutes
 
0.5 mark for each axis with correct labels and units. (1 mark for x and y axes)
1
0.5 mark for indicating the time for incipient plasmolysis and full turgor on the x axis
1
0.5 mark for labelling the curves
(5)
Transport Structures in Vascular
Plants
 Transport Structures in Vascular
Plants
• Root
– support
– Transport ：
Epidermis
Cortex
Endodermis
Stele
（ incl. xylem
and phloem)
 Structural adaptation of root hair cells

Feature Function
Long extension increase surface area : free contact with soil water and ions
Thin/ Fine/ Slender penetrates soil particles for soil water contact
Large vacuole give osmotic control
Thin/ Unthickened cell wall for absorption of water and mineral salts
Absence of cuticle for absorption of water and mineral salts
Abundance of mitochondria for absorption of mineral salts by active transport
 Herbaceous Dicot Stem Structure

• vascular bundles in a ring near the epidermis

• Vascular bundle includes a vascular cambium
– secondary growth
Epidermis
Collenchyma: living elongated
cells with cell walls thickened at
corners for extra support, able to
stretch (growth)
 vacuole

thin cell wall

Cortex: Parenchyma cells with thin walled cells

Sclerenchyma:
dead when mature,
with thick deposits
of lignin, e.g.
sclereids and fibres
 Structure/Function: Xylem
• Three types of xylem cells
– Tracheids
• elongate with tapered ends where
cells are connected into long tubes
• pits allow communication
between tracheids
– Vessel Elements
• highly specialized for transport
• many open-ended
• form vessels
– Rays
• lateral transport
Vessel Elements
Pulling Xylem Sap: The Transpiration-
Cohesion-Tension Mechanism
• Water is pulled upward by negative
pressure in the xylem
Transpirational Pull
 Cohesion and Adhesion in the
Ascent of Xylem Sap
• 1 A negative pressure ( tension ) is created on the surface of the film of water coating
mesophyll cells when evaporation occurs from its surface. (This occurs during
transpiration).
• 2 This tension pulls on the interconnected chain of water molecules within the xylem
which extends from the leaf through the stem to the tips of the root.
• 3 The water molecules in the chain are held together by hydrogen bonds which exist
between neighbouring water molecules. ( cohesion)
• 4 The chain of molecules is prevented from being pulled down because each water
molecule in the chain is attracted to the walls of the xylem by hydropyllic attraction
between water and the cellulose in the cell walls. (Adhesion)
• 5 Hence the water column which is held together by cohesion and prevented from
lowering by adhesion is pulled up by the tension generated from above by
transpiration.
• 6 Solutes which are dissolved in the water also are pulled up within the xylem.
 • Ascent of xylem sap
Xylem sap
Outside air  = –100.0 MPa
Mesophyll cells
Stoma
Leaf  (air spaces)= –7.0 MPa Water molecule
Transpiration
Atmosphere
Leaf  (cell walls)= –1.0 MPa

Water potential gradient

Xylem
cells
Cell wall

Adhesion

Trunk xylem = – 0.8 MPa Cohesion, by hydrogen

Cohesion bonding
and adhesion
in the xylem
Water molecule

Root xylem = – 0.6 MPa Root hair

Soil = – 0.3 MPa Soil particle

Water uptake Water
from soil
Root Pressure
When a plant is carefully severed close to the base of the
stem, sap oozes from the stump. The fluid comes out under
pressure which is called root pressure.

Root pressure is created by the osmotic pressure of xylem

sap which is, in turn, created by dissolved minerals and
sugars that have been actively transported into the apoplast
of the stele.

Although root pressure may play a significant role

in water transport in certain species or at certain
times, most plants meet their needs by
transpiration-pull.
2005 Paper IIB Q.5
2009 Paper II
Pathways of water absorption in roots
Casparian strip
Endodermal cell
Pathway along apoplast

Pathway through
4 5
symplast

Casparian strip
Plasma
membrane (
Apoplastic
1
route
2 Vessels
3 5
4 (xylem)
Symplastic Root
route hair
Vascular
Epidermis Cortex
Endodermis cylinder
The Pathway of Water in a plant root:
1) Apoplastic
2) Symplastic 3) Vacuolar
Soil water enters the root at the root hairs -
extensions of epidermal cells. It appears that water
then travels in both:

•in the nonliving parts of the root - called the

apoplast - that is, in the spaces between the cells
and in the cells walls themselves. This water has
not crossed a plasma membrane.

•the cytoplasm of root cells - called the symplast

- that is, it crosses the plasma membrane and then
passes from cell to cell through plasmodesmata.
However, the inner boundary of the cortex, the
endodermis, is impervious to water because of a
band of suberized matrix called the casparian
strip. Therefore, to enter the stele, apoplastic
water must enter the symplasm of the endodermal
cells. From here it can pass by plasmodesmata
into the cells of the stele.

Once inside the stele, water is again free to move

between cells as well as through them. In young
roots, water enters directly into the xylem vessels
and/or tracheids. These are nonliving conduits so
are part of the apoplast.
Mechanism of Water Absorption

Passive Absorption
Passive absorption is by osmosis. Passive absorption takes
place along the concentration gradient - when the
concentration of cell sap is higher than that of soil water.

Water is absorbed when transpiration rate is high or soil is

dry. Due to high transpiration rate, water deficit is created
in transpiring cells. Rapid transpiration removes water and
reduces turgor pressure in living cells of root.

The suction force thus developed is transmitted to root

xylem. It pulls water from surrounding root cells to make up
water deficit.
uptake of Minerals
Minerals enter the root by active transport (pumping
against the concentration gradient using ATP) into
the symplast of epidermal cells and move toward and
into the stele through the plasmodesmata connecting
the cells.

They return to the apoplast from the cells of the pericycle

through specialized transmembrane channels.

Once in the xylem, water with the minerals that have

been deposited in it move up in the vessels and tracheids.
At any level, the water can leave the xylem and pass
laterally to supply the needs of other tissues.
 Structural adaptation of xylem for transport

Cells types Features Adaptation

one elongated cell on top of another
forming vertical column
 hollow tubes result and facilitate mass/free flow of water
end wall and cell content lost 
cell dead

 to prevent collapse/pressure change due to -ve tension

cell wall lignified and strengthened develop during active transpiration
Xylem vessel  to give mechanical support
 

 allow inter-communication between cells

pits  vessels for free passage of water
 prevent air lock in transpiration stream

hex/octagonal shape  close packing for strength

elongated hollow cell, imperforated
with pit-closing membrane present in  water flow from cell to cell through pits
Tracheid the region of pits

cell wall lignified and strengthened  to give mechanical support

long cell with cell wall thickened and

Fibre  to give mechanical support
lignified
Parenchyma cell cell wall may or may not be lignified   as storage cell. 
Vessel Elements
Transpiration is thought to
occur because the plant
requires CO2 from the
surrounding air for
photosynthesis. For most
plants, stomata open in the
morning after the plant has
been exposed to the sun
long enough to use up much
of the CO2 inside the leaf,
and stomata close at night.
The loss of water is
incidental to the need for
uptake of CO2.
 Woody stems and mature roots are
sheathed in layers of dead cork cells
impregnated with suberin - a waxy,
waterproof (and airproof)
substance. So cork is as impervious
to oxygen and carbon dioxide as it is
to water.

However, the cork of both mature

roots and woody stems is perforated
by nonsuberized pores called
Lenticular & lenticels. These enable oxygen to
cuticular transpiration reach the intercellular spaces of the
interior tissues and carbon dioxide to
be released to the atmosphere.
Guard Cells

Leaf surfaces are dotted with millions of

stomata such as this one. This stoma is
lined by two guard cells that control its
aperture. Because control requires
movement, and movement requires
energy, these cells contain numerous
mitochondria and chloroplasts. Thus
they are the only cells in the epidermis
that are green.
The opening or closing of the stomata is a
result of the changes in the turgor pressure of
the guard cells. The inner wall of each guard
cell is thick and elastic. When turgor
develops within the two guard cells flanking
each stoma, the thin outer walls bulge out
and force the inner walls into a crescent
shape. This opens the stoma.

When the guard cells lose turgor, the elastic

inner walls regain their original shape and
the stoma closes.
(OS) The stomata will open when cells are turgid. The reason for the
changes in turgor pressure is the movement of water from subsidiary
cells into the guard cells. This is a passive movement caused by
changes in water potential created by a K+ pump (active transport).
 Transpiration
• Transpiration is the loss of water from a plant by evaporation

• Water can only evaporate from the plant if the water potential is lower in the
air surrounding the plant

• Most transpiration occurs via the leaves

• Most of this transpiration is via the stomata.

58
 Classical starch theory
• Morning CO2 concentration decreases and
• And pH of guard cell increases which
activates Phosphorylase.
• Starch + n H3PO4 Glucose1PGlucose 6-
P + H2OGlucose+ H3PO4
• Enzymes- Phosphorylase, Phosphomutase, Phosphatase.
• Glucose increase osmotic conc. Of guard
cell and initiate absorption of water. Which
leads to swelling of guard cell and thus its
opens.
Mechanism of Stomatal opening
• Malate or K+ ion pump Hypothesis (modern
theory)
• Levitt (1974) propose pH of the guard cell
rise due to active H+ uptake by guard cell
chloroplast or mitochondria, CO2
assimilation by mesophyll and guard cell.
• Starch--- Hexose 6-P-------Phosphoric acid-
Phosphoenol pyruvate
• PEPcarboxylase convert
• PEP + CO2 Malate2-
Structure of STOMATA
 How does transpiration occur?
• Due to endosmosis of water turgor pressure of
guard cells increase.

• Outer wall stretches and bulges out.

• Inner wall gets pulled apart.

• Stomata open.
 Stomatal movement – how?
• Movement of stomata can be explained by active
potassium ion transport mechanism.
• It was proposed by FUJINO and later explained
by LEVITT.
• According to this theory opening of stomata is an
active process whereas closing is a passive
process.
In day
time
Phosphoenol pyruvate

Malic acid
Dissociation of malic acid into malate ion and
proton
Efflux of proton and influx of potassium ion

Potassium and malate ion in guard cell

Osmotic pressure of GC increases

Endosmosis in guard cells

STOMATA OPEN
In night
time
CO2 accumulates in GC

pH decreases and ABA becomes active

Potassium ion is transported back out

Formation of starch

Osmotic pressure of GC decreases

Exosmosis in guard cells

Guard cells become active

STOMATA CLOSED
 Antitranspirants
• During drought conditions the plant shows
wilting.
• In arid and unirrigated areas crop plant
cannot grown because of excessive
transpiration and reduced water availability.
Substances that reduces the rate of
transpiration are called antitranspirants.
• Phenyl mercuric acetate, Abscisic acid,
silicon emulsion, colorless plastic resins.
Distribution and Number of Stomata
More stomata are found on the ventral
leaf surface, away from sunlight and
wind. Some plants have no stomata at all
on the dorsal leaf surface.
Transpiration does serve three useful
functions:
•It provides the force for lifting the
water up the stems.
•It cools the leaves.
•It helps absorb soil water.
 Measuring rate of
transpiration

A decrease in mass of the apparatus is

a measure of the mass of water lost
from the leafy shoot by transpiration
under varying environmental
conditions.
Factors Affecting the Rate of Transpiration

1. Air Temperature
Leaves receiving direct
sunlight absorb about 80% of
the radiant energy falling on
them. Part of this is changed
to heat, raising the internal
temperature of the leaf. Part of
this heat is used in vaporizing
water and a small part is used
 Environmental factors
2. Relative humidity of the
atmosphere
The drier the air around the plant, the
greater the transpiration rate.

The intercellular spaces and the cells

of a turgid leaf are usually almost
saturated with water. In this case a
"steep" diffusion gradient would exist
between the leaf and the atmosphere.
The
Increased rate =ofIncreased
vapour transpiration would
diffusion gradient =
therefore
Increased be high.
transpiration
3. Light Intensity
Illumination stimulates the opening of
the stomata and increases
transpiration.

4. Wind/Air Movement
Removes the mantle of moist air
surrounding the leaf. If dry air is
blowing across the leaf the
diffusion gradient will be steep
and rapid transpiration results.
5. Water Supply
Any soil condition affecting the
absorption of water by the roots will
affect the rate of transpiration.
If the rate of water loss
(transpiration) exceeds the rate of
absorption by the roots, wilting will
occur.
H.W. p. 205, Q.5
Hand in this Wednesday
Patterns of Phloem Translocation
•Sugar transport in the Phloem is NOT regulated by
gravity. It is Bi-directional. There is a general
pattern of carbohydrate redistribution from Sources
towards Sinks.
•Sources are carbohydrate exporting structures like
mature, photosynthetic leaves. Storage organs like
the potato tuber can also be Sources.
•Sinks are carbohydrate importing structures like
immature, nonphotosynthetic leaves.

•Storage organs can be Sinks or Sources. The storage

root is a Sink during its first year of existence.
However, it becomes a source during its second year
when it provides the nutrients for flower production.
2003 Paper IIB 4 (a)
Actively growing Shoot & Root Tips are powerful
Sinks. Developing Ovules & Fruits are also strong
Sinks.
 Structure/Function: Phloem

• Sieve tubes are made of sieve elements

– connected by sieve plates
– intact cytoplasm, no nucleus
• Companion cells
Mass flow: Radiocarbon as a tracer
 Mouth part as micropipette

• Aphids, such as greenfly, have specialised

mouthparts called stylets, which they use to
penetrate phloem tubes and suck the sugary
sap. If the aphids are anaesthetised with
carbon dioxide and cut off, the stylet remains
in the phloem so pure phloem sap can be
collected through the stylet for analysis. This
surprising technique is more accurate than a
human with a syringe and the aphid’s
enzymes ensure that the stylet doesn’t get
blocked. Phloem is confirmed to transport food.
 The Mass Flow Hypothesis
phloem
sink

source

A gradient of pressure potential exists between the

source & the sink with phloem linking them and as
result liquid flows from the leaves to other tissues
along the sieve tube elements.
Mechanisms of translocation in phloem Mass
Flow (pressure flow) Hypothesis - widely accepted
At the source (leaves):
Photosynthesizing cells in leaves make sucrose
 water potential decreases
 water enters cells from xylem creating high
pressure potential
At the sink (root):
sucrose either respired or stored as starch
 water potential increases
 low pressure potential
2005 Paper II B Q.5
2008 Paper II B Q.4 (b)
Evidences supporting the mass flow theory:
[Link] is a flow of solution from phloem when it is
cut or punctured by the stylet of an aphid
[Link] of concentration gradients:
high concentration of sucrose in leaves
but low concentration in root
[Link] researchers have observed mass flow in
microscopic sections of living sieve elements
• Criticism:
• Sieve plates could be a series of barriers
impeding flow since the process is passive
after the initial process.

• Suggested reason for sieve plate function:

seal off damaged sieve tube elements with
callose deposits across the pores.
• Summary of mass flow hypothesis:
• 1) Sucrose made in leaves/by photosynthesis/from
breakdown of storage compounds;
• 2) (Sucrose) actively transported into phloem;
• 3) by companion/transfer cells:
• 4) water potential decreases;
• 5) water enters by osmosis;
• 6) hydrostatic pressure increases;
• 7) In growing areas/sinks /storage areas sucrose is
removed;