Chapter 6

Plant Water Relation

• Water -‘elixir of life’ constituting about 90 to 95% of most plant
cells and tissues.
• It maintains turgidity and shape of plant cells and organs.

Properties of water:
1. A significant molecule that connects physical world with
biological processes.
2. Liquid at room temperature.
3. It is the best solvent for most of the solutes and hence the best
aqueous medium for all biochemical reactions.
4. In pure form it is inert with neutral pH.
5. The best transporting medium for dissolved minerals and food
molecules. An essential raw material for photosynthesis.
6. It has high specific heat, high heat of vaporization and high heat
of fusion, thus acts as thermal buffer. These various properties
are due to hydrogen bonds between the water molecules.
7. Water molecules have good adhesive and cohesive forces of
attraction and high surface tension.

Water absorbing organ: Root :

o In terestrial plants, roots absorb water and minerals from the
soil.
o A typical root has unicellular, epidermal hair in the region of
absorption.
o Root hair is a colourless, unbranched, delicate cytoplasmic
extension of epiblema cell.
o It has a plasma membrane and thin two layered cell wall, outer
layer of pectin and inner of cellulose.
o Cell wall is freely permeable but plasma membrane is
selectively permeable.
Water available to roots for absorption:
The microenvironment surrounding the root from where plants
absorb water is called rhizosphere.
Water in the soil occurs as-
• Gravitational water (that percolates deep due to gravity)
• Hygroscopic water (water adsorbed by soil particles)
• Combined water (water present in the form of hydrated oxides of
silicon, aluminium, etc.)
• Capillary water (water held in pores between the soil particles).
Of these forms only the capillary water is available for absorption.

Absorption of water by roots from soil:

Root hair absorbs water by Imbibition, diffusion and osmosis.
a. Imbibition:
• The swelling up of hydrophillic colloids due to adsorption of water
is called Imbibition.
• Imbibant - Substance that adsorbs water.
• Imbibate - Water/ liquid that gets imbibed.
• The root hair cell wall is made up of hydrophilic colloids (pectic
compounds and cellulose) which adsorb water.
• Imbibition is important in seed germination, swelling up of dried
raisins, kneading of flour etc.

b. Diffusion: Fig 6.2

• It is the movement of ions/ atoms/ molecules of a substance from
the region of their higher concentration to the region of their lower
concentration due to the kinetic energy of the molecules.
• Water diffuses into the cell through a freely permeable cell wall.
• Diffusion results in the diffusion pressure (D.P.) which is directly
proportional to the number of diffusing particles.
• D.P.of pure solvent > D.P. of solvent in a solution.
• The difference in the D.P.values of pure solvent and the solvent
in a solution is called Diffusion Pressure Deficit (DPD) or
Suction Pressure (SP) or Water potential.
• DPD=The thirst of a cell with which it absorbs water from the
surroundings.
• Diffusion is significant in plants in the absorption of water,
minerals, conduction of water against the gravity, exchange of
gases and transport and distribution of food.

c. Osmosis:
 It is a special type of diffusion of solvent through a
semipermeable membrane.
 The cytoplasm of root hair cell is more concentrated (due to
dissolved minerals, sugars, etc.) than outside the cell which
results in the flow of solvent into cytoplasm through the semi
permeable plasma membrane.
 With respect to the concentration and osmotic migration,
solutions may be of 3 types
i. Hypotonic with low osmotic concentration,
ii. Hypertonic with high osmotic concentration and
iii. Isotonic neither gain nor loss of water in an osmotic
system.
  Osmosis may be Exo-osmosis and Endo-osmosis.
 Exo-osmosis is the migration of solvent from the cell outside
making the cell flaccid.
 Endo-osmosis is the migration of the solvent into the cell
making the cell turgid.
 Turgidity increases the turgor pressure (T. P.) of the cell
which is the pressure exerted by turgid cell sap on the cell
membrane and cell wall.
 Cell wall exerts a counter pressure on the cell sap called Wall
pressure (W. P.).
 In a fully turgid cell, DPD is zero.
 In a fully turgid cell, T. P. = W. P. but operating in opposite
direction.

Osmotic pressure (O. P.):

The pressure exerted due to osmosis is osmotic pressure.
Which is equivalent to the pressure which must be exerted
upon it to prevent flow of solvent across a semipermeable
membrane.
Therefore, D. P. D. = O. P. - T. P.
As T. P. = W. P., D. P. D. = O. P. - W. P.
In a flaccid cell, T. P. is zero therefore, DPD = OP
In a turgid cell, DPD is zero ∴ TP = OP

Facilitated diffusion:
• It is the passive absorption of solutes mediated by a carrier.
• Lipid soluble particles that are can easily diffuse through
lipoproteinous cell membrane. However, the diffusion of
hydrophilic solutes has to be facilitated.
• Special membrane proteins called aquaporins and ion- channels
provide sites for facilitated diffusion.
• These proteins help move substances across membranes along
the concentration gradient without the expenditure of energy.
Water Potential (ψ):
• Free energy per molecule in a chemical system, is called its
chemical potential and Chemical potential of water is called
water potential.
• It is represented by psi (ψ) has a negative value.
• Water potential of pure water is always zero. Addition of any
solute in it, decreases its psi (ψ) value. Therefore, it has negative
value.
• Water potential of protoplasm is equal but opposite in sign to
DPD.
• It is measured is in bars/ pascals/ atmospheres.
• Water always flows from high water potential to low water
potential. Difference between water potential of the adjacent cells
decides movement of water through plasmodesmata across the
cells.

Factors affecting water absorption:

 Presence of capillary water,
 temperature (between 200 to 300C),
 concentration of solutes in soil water,
 aeration of the soil and
 transpiration.

Plasmolysis:
 It is the exo-osmosis in a living cell when placed in hypertonic
solution.
 The protoplast of cell shrinks and the cell becomes flaccid.
 When placed in hypotonic solution, the cell becomes turgid by
endo-osmosis. This is called deplasmolysis.
 A plasmolysed cell - T. P. is zero.
 A fully turgid cell T. P. = O. P. hence, DPD is zero.
Path of water across the root (i.e. from
epiblema to xylem)
• The root hair cell gains water by imbibition, diffusion and osmosis
and becomes turgid.
• It has now high T.P., but low DPD. The adjacent cortical cell with
high O.P., has more DPD value.
• As a result, water moves from the root hair cell to the cortical cell.
Root hair cell is then ready to absorb water from the soil.
• Water from the turgid cortical cell moves to inner cortical cell and
the process continues.
• There is a gradient of suction pressure (DPD) developed from
cells of epiblema to the cortex of the root. The water further moves
into the cell of pericycle via passage cells of endodermis and
finally into the protoxylem.
• The hydrostatic pressure is developed in living cells of root (root
pressure) due to which, water from pericycle is forced into the
xylem and then upwards against the gravity.
• Water moves across the root by apoplast and symplast
pathways.
• Apoplast pathway - movement of water through the cell wall and
intercellular space
• Symplast pathway - movement of water through Plasmodesmata.
Mechanism of absorption of water :
a. Passive absorption:
• The driving force - transpiration pull
• No expenditure of energy.
• Water moves along the concentration gradient
• Occurs during daytime when transpiration is active.
b. Active absorption:
• Water is absorbed due to activity of roots.
• The driving force - the root pressure, in the living cells of root.
• Occurs usually at night when transpiration stops.
• It is against the DPD gradient.
• Energy is used which is generated through the respiration.
• Active absorption may be osmotic and non-osmotic.
• In osmotic absorption water is absorbed from soil into xylem of
the root along the osmotic gradient. Energy is required to create
osmotic conditions.
• In non-osmotic absorption water is absorbed from soil against
the concentration gradient. It requires energy released during
respiration, directly. Poor supply of oxygen, low temperature and
metabolic inhibitors retard water absorption as they decrease the
rate of respiration.

Translocation of water (Ascent of sap)

• It is the upward transport of water with dissolved minerals from
root to other aerial parts like stem and leaves, against the gravity.
• It occurs through the lumen of tracheids and vessels, in all
vascular plants.
Following three theories explain the mechanism of
ascent of sap:
a. Root Pressure Theory (Vital Theory):
• Due to continuous water absorption by roots, a hydrostatic
pressure is developed in the living cells of cortex of root called
root pressure.
• Water along with dissolved minerals is forced into xylem and also
conducted upwards against the gravity It is due to root pressure.
• Root pressure development is an active process and is
influenced by factors like oxygen, moisture, temperature of soil,
salts,etc.
• The value of root pressure is +1 to +2 bars.
• The theory however is not applicable to plants taller than 20
meters.
• The theory fails to explain the ascent of sap in the absence of
root system, zero root pressure value in taller gymnosperm trees,
absence of root pressure in actively transpiring plants,etc.

b. Capillarity theory (physical force theory):

• It suggests the involvement of physical forces and dead cells in
the ascent of sap.
• The lumen of tracheids and vessels function as capillaries.
• Water column is formed due to cohesive and adhesive forces of
water and xylem wall as a result water is raised upwards against
the gravity.
• However, tracheids with closed end-walls, presence of a barrier
of root cortex between xylem and soil water and presence of wider
xylem vessels than tracheids in tall plants do not support this
theory.

c. Cohesion- tension theory (Transpiration pull

theory): (Dixon and Joly)
• It is the widely accepted theory that explains the ascent of sap in
plants.
• It is based on the concepts of
I. Cohesion and adhesion, and
II. Transpiration pull.
Cohesive force of attraction exists between water molecules
while there is adhesive force of attraction between water
molecules and lignified wall of lumen of xylem vessel. These two
forces develop a continuous water column in the xylem.
• Due to continuous transpiration through stomata of the leaf, the
D.P.D. of mesophyll cells increases. They withdraw water from
xylem in the leaf.
This results in a tension called transpiration pull.
• Due to the transpiration pull the water column pulled upwards
passively against the gravity. This pull is responsible for the ascent
of sap.
• However, occasional breaks in the water column due to gas
bubbles, ascent of sap in absence of transpiration and in absence
of leaves in deciduous trees, etc. do not support this theory.

Transport of mineral ions :

• Minerals play an important role in plants and they are absorbed
in the dissolved form through roots.
• They are transported upwards along with xylem sap. Mineral ions
are required in the meristems, young leaves, developing flowers,
fruits, seeds and storage organs.
• They enter the cells by active absorption.

Transport of food:
• The leaves where food is synthesized constitute the source and
while part where it is utilized, is called sink.
• The food is transported from leaves to the non-green parts like
root and stem.
• Food is always translocated in the form of sucrose along the
concentration gradient from source to sink.
• Food is translocated longitudinally through sieve tubes of
phloem. It is translocated in downward from leaves (source) to
stem and root (sink). It also occurs in upward direction during
germination of seed, bulbils, corm and from leaves to the growing
points.
• The lateral translocation occurs via medullary rays in the stem.
• From phloem to pith - Radial translocation
• From phloem to cortex - Tangential translocation
• Thus, the transport of food through phloem is bidirectional.
• Phloem sap contains mainly water, sucrose, amino acids and
hormones.

Mechanism of sugar transport through

phloem:
(Munch’s pressure flow theory or mass flow hypothesis)
• The photosynthetic production of glucose increases the osmotic
concentration of the cell. As a result, the cell becomes turgid due
to endo- osmosis.
• Due to increase in turgor pressure, sugar from photosynthetic cell
is forced into the sieve tube of the vein which is called Loading of
Vein.
• At the sink end, the cells utilize sugar or polymerizes excess
sugar into the starch lowering its osmotic concentration.
• Exo-osmosis occurs, thereby decreasing the turgidity of cell. Due
to lowered turgor pressure, food is translocated along the
concentration gradient, passively. This is Vein unloading. At the
sink end sugar is used and excess water exudes into the xylem.
• This theory, however, does not explain bidirectional transport of
food.

Transpiration :
 The loss of water in the form of liquid through water stomata
or hydathodes is called guttation.
 The loss of water in the form of vapour through cuticle,
stomata and lenticels is called transpiration.
1. Cuticular transpiration:(8-10%) Loss of water through a
layer of waxy substance- cutin, called cuticle
2. Lenticular transpiration:(0.1-1.0%) Fig 6.7 Lenticels are
small raised structures composed of porous tissue with
loosely arranged cells. Lenticels are present in bark of old
stem and pericarp of woody fruits but are absent in leaves.
3. Stomatal transpiration:(90 to 93%) Fig 6.8 a,b
 Stomata are minute apertures in the epidermis of young stem
and leaves.
 Depending upon distribution of stomata on leaves, leaves
may be
epistomatic- on upper epidermis (Hydrophytes-e.g. Lotus),
hypostomatic- on lower epidermis (Xerophytes- e.g. Nerium)
amphistomatic-on both surfaces (Mesophytes- e.g. Grass).
Structure of stomatal apparatus :
• A stoma consists of two guard cells, stoma and accessory
cells(subsidiary cells).
Guard cells are kidney-shaped in dicot plants and dumbbell-
shaped in monocot plants(grasses). Fig 6.9
• Guard cells are living, nucleated cells with inner thick and
inelastic wall and outer thin and elastic wall is.
Opening and Closing of Stoma :
• Guard cells turgid - Stoma opens .Guard cells flaccid – Stoma
closes.
• According to starch-sugar inter-conversion theory (Steward),
• During day time, startch is converted into sugar by the enzyme
phosphorylase. The osmotic potential of guard cells increases.
• Entry of water there by gaurd cells and stoma widens.
• Durind night time, the reverse reaction occurs causing the
stomatal closure.
• According to theory of proton transport (Levitt), stomatal
movements are controlled by the transport of protons H+ and K+
ions.
• During daytime, starch is converted into malic acid.
• Malic acid dissociates to form Malate and protons.
• Protons are transported to subsidiary cells and K+ ions are
imported from them. Potassium malate is formed that increases
osmolarity and causes endosmosis. Uptake of K+ ions is always
accompanied with Cl¯ ions.
• At night, abscissic acid changes the permeability of guard cells
thereby preventing the uptake of K+ and Cl- ions.
• As a result guard cells become lose water and become flaccid
causing stomatal closure.
Significance of Transpiration:
 Excessive transpiration leads to wilting and injury in the plant.
Stoma must remain open for the gaseous exchange.
 It removes excess of water and helps in the ascent of sap.
 It maintains turgor of the cells and imparts a cooling effect.
Hence, transpiration is ‘A necessary evil ’.