W ATER R ELATIO N S O F

Importance
of
water to plants:
P
LA
N
TS
Water is the most vital substance necessary to carry out the

activities of life. It is of following importance.

Component of protoplasm: It is the essential and major
component with out which the cell becomes in active or killed.
Universal solvent: It is the excellent solvent in the cytoplasm
that carries various cellular reactions.
Transport: it is involved in the distribution of dissolved
substances from one cell to other in turn one region to the
other.
Break down of Chemical substances: The break down of
chemical substances in cells requires water.
Absorption: plants absorb nutrients from soil in the solution
form.
Turgidity: Mobility of Gametes, Dispersal of spores, pollination,
dispersal of fruits takes place due to turgidity.
Support: it gives support (rigidity) to aquatic plants.
Source of Hydrogen: e.g. Photosynthesis.

The water absorption and transportation system includes Imbibition, Diffusion, Osmosis and Plasmolysis.

Imbibition
It is the process of movement of water towards the Hydrophilic
region (end) of the substance is called Imbibition.
Example: Putting of Pea seeds in water, during which they get
soaked and swell in size.

Factors affecting Imbibition:

Effect of temperature: With increase in temperature the
imbibition also increases.
Effect of Concentration: The rate of imbibition decreases with
increase in the concentration of the solute.
Effect of pH: Affected by Alkalinity and acidity. Hence require
definite pH.

Significance:
Initially plant roots absorb water by Imbibition.
It is the first step in the germination of seeds.

Diffusion:
The process of movement of particles from the region of higher
concentration to the region of lower concentration, until the
achievement of equilibrium is termed as Diffusion.
During Diffusion the molecules are evenly distributed.
Examples: Opening of Scent bottle, LPG leakage, H 2S gas spreading.

Factors affecting Diffusion:

Temperature: With increase in temperature the Diffusion also
increases.
Concentration: The rate of Diffusion decreases with increase in the
concentration of the solute.
Pressure: The rate of Diffusion increases with increase in the
Pressure.
Vacuum: The rate of Diffusion decreases in the vacuum.
Significance:
It permits the material exchange between the cells, cells and
environment. E.g. Salt/ gas exchange.
It takes part in the transpiration.
It plays little role in translocation of substances.

Osmosis
Nollet (1748) first demonstrated the osmosis.
Osmosis may be defined as the movement of
molecules from the region of higher concentration to
the region of lower concentration through semi
permeable membrane or selectively permeable
membrane.
OR
If diffusion takes place through the semi permeable
membrane.
Endosomosis: the movement of solvent molecules

takes place from outside (higher concentration) to

inside (lower concentration)
Exosmosis: the movement of solvent molecules
takes place from inside (higher concentration) to

 Solvent: the substance in which other substances are dissolved.

Solute: the substance that is dissolved in solvent.
Solution: the mixture of solute and solvent.
Semi permeable membrane: the membranous structure that

allows only the solvent and some other substances to pass through it.
Hypertonic solution: the solution with more amount of solute and

less amount of solvent is regarded as strong solution.

Hypotonic solution: the solution with fewer amounts of solute and

more amount of solvent is regarded as weaker solution.

Isotonic

solution: When the solution is kept in (similar

concentration) another media if there is movement of solvent
molecules then it is said to be the isotonic solution OR the equal
concentration of two solutions when mixed together in which there is
no movement of molecules is said to be the isotonic solution.

 Plasmolysis: the shrinkage of cytoplasm

away from the cell wall due to exosmosis is

called Plasmolysis. During plasmolysis the
cells undergo flaccid condition.
Example: plant cell kept in hypertonic solution

shrinkage takes place.

Deplasmolysis: The swelling of cytoplasm

away from the cell wall due to endosmosis is

called Deplasmolysis. During deplasmolysis
the cells undergo turgid condition.
Example: plant cell kept in hypotonic solution

swelling occurs.

Turgor pressure and wall pressure

The pressure created by the solvent molecules (water)
during endosmosis in a cell is turgor pressure.
When plant cell is put in water endosmosis takes place
and the cell becomes turgid and pressure is
developed.
As the turgor pressure increases, it exerts a pressure
on the cell wall. So the cell wall resists with an equal
amount of pressure and it is called wall pressure.
The free energy of solvent molecules is altered by the
addition of solute or temperature or pressure.
The addition of solute decreases the free energy. The
increase in the temperature and pressure increase the
free energy.
The solvent molecules move from the region higher
free energy to the lower free energy. There fore water
movement occurs on the pressure gradient.

Water potential ( w)
Difference between the free energy of water

molecules in pure water and the free energy water in

every solution.
Represented by w
Measured in terms of bars or Pascals.
The value of pure water is always is zero and it is
considered to be the highest water potential.
If little solute is added to this water, the water
potential is lowered and it is less than zero. There
fore it is always ve value.
This means water potential involves the flow of
water molecules from the region of higher water
potential to the region of lower water potential. I.e.
less negative side to more negative side.

Osmotic potential ( s)
The amount by which the water potential is
reduced due to the addition of solute molecules
to water.
Represented by s
This is always zero.

Pressure potential ( p)
It is developed due to turgor pressure and wall
pressure. It is always positive and increases the
water potential of the cell sap.
The relation ship between the Water potential,
Osmotic potential and Pressure potential is given
by the equation
w = s + p

 Suppose there is a fully turgid cell having osmotic

solution of -5 bars and pressure of 5 bars then the

water potential is zero.
w = s + p
w = -5 + 5
w = 0

On the contrary flaccid cell has no turgor pressure

and so there no pressure potential (zero) suppose the

osmotic potential of cell sap is 5 bars the water
potential of the cell is 5 bars the relation is given by
w = s + p
w = -5 + 0
w = -5

Absorption of water
The plants absorb water and minerals from soil through
the root hairs of the root.
The structure of Root and Root hair
The root has the following structures
Root cap: tip of the root protects the root from injury.
Meristematic region: region of cell division and
growth present above the root cap
Region of elongation: above the Meristematic region
where cells show the elongation process.
Region of root hairs: Above the elongation region
1mm in thickness bear up to 2 million minute root hairs.
Root hairs: some of the epiblema cells are elongated
and specialized to form the root hairs. They are short
lived and are hydrophilic in nature as they contain
Pectin and cellulose.

Sources of water for plants

The primary source of water for plants is the soil
water.
The chief source of water to the soil is the
rainwater.
The soil water occurs in different forms in the soil.
The total amount of water present in soil is called
Holard.
The water available to the plants is called
Chresard.
The water not available to plants is Echard.

Echard includes
Runway water: that flows down the slopes to the other
regions.
Gravitational water: water that percolates down and
deep in to the soil. It is beyond the plants reach.
Hygroscopic water: water held by soil particles due to
adhesion.
Chemically combined water: water held by minerals
like Iron, Al, Si, etc.
Chresard includes
Capillary water: the water is held between the soil
particles due to surface tension. It depends upon the
type of soil. It is only the easily available water for plants.
Hence in Agriculture the drip irrigation is practiced. In
which the water remains around the plants in the soil.

Region of Absorption of water in plants:

Water absorption in higher plants takes place
through the root system.
In lower and aquatic plants it is absorbed
through the surface.
How ever the water absorbed is utilized for
the metabolic activities of life.
The maximum amount of water absorption
takes place in the region of root hairs, in this
region cells are vacuolated, xylem is well
differentiated and some of the cells of the
epiblema formed the root hairs.

Entry of water from soil in to Xylem of root

Water absorption occurs in the root hair, then passed in to

the cells of the cortex, then in to the passage cells of the

endodermis the in to the Xylem.
There are three possible pathways for the entry of water

from soil in to the plant

Apoplast pathway: the water moves through the network

of nonliving material such as the cell wall and middle lamella.

Symplast pathway: the water moves through the network

of living material such as the cytoplasm and plasmodesmata.

Vacuolar pathway: the water moves through number of

membranes, cytoplasm and vacuoles by osmosis.

Mechanism of Absorption of Water

Two types of Mechanisms is observed in the absorption of
water in plants namely - Passive absorption and Active
absorption.
Passive absorption: in this type of water absorption
involves no expenditure of energy. It is of two types
Water potential Gradient:
The capillary water -water contains solutes (organic and

inorganic) and remains as Soil solution (Dilute solution).

The water potential of soil solution is less than the pure
water i.e. less than Zero.
Where the water in the cell sap of the cells contains more

solutes (organic and inorganic) remains as concentrated

solution. There fore the water potential of cell is lesser
than the Soil solution.
The two regions are separated by plasma membrane.

Naturally it results in endosmosis. i.e. the movement of

water takes place from the region of higher potential to

Transpiration Pull:
Transpiration results in the decrease in the water
potential of the cells of mesophyll of leaves, which
results in the transpiration tension.
Transpiration tension develops a suction force that
sucks water from the lumen of Xylem vessels, which
is termed as transpiration pull.
Due to transpiration pull the water potential in stelar
region is decreased.
Naturally soil water potential is more that moves in
to the less water potential zone
Non-osmotic active absorption:
It is observed in the epiblema of certain plants and
Animal cells lining in the Nephron of kidney,
Intestine etc
The absorption of water occurs by means of certain

Osmotic active absorption:

This concept was explained according to Craft and Broyer (1938).
According to this theory the root is acting as Osmoscope. The
tissues of the cortex have high water potential since water could
enter in to these tissues easily by imbibition / capillary action /
Cell wall material.
The tissues in the stele has lower potential because of the
continuous action of apical meristem in absorption mineral
solution and subsequent pumping of excess of solutes in to the
stelar region. Both the activities require ATP.
Factors affecting the absorption of water in plants:
Availability of water:
Soil temperature:
Concentration of the solution:
Aeration of Soil:
Metabolism of plant:
Root system:
Rate of transpiration:

Ascent of Sap
Ascent of Sap may be defined as the upward movement of water
and minerals from the region of availability (roots) to the region of
utilization (shoot) through the conducting path (stem).
The water absorbed from the soil in to the plant reaches the xylem
through root hairs, cortex, and endodermis.
From the xylem it moves in upward direction and reaches the
mesophyll region of leaves where little amount of water is utilized
for metabolic activities and greater amount is loosed by
transpiration.
Evidences to show the involvement of Xylem in Ascent of
Sap
The rooted Balsam plant is selected for study. The roots of Balsam
plant is immersed in dilute colored solution (Eosin / Safranin) set
up is left aside for 30-60 minutes in Sun light.
A red line is seen moving in the upward direction.
When T.S. of that stem is taken and observed under microscope
the region of xylem is colored which indicates the xylem is
involved in the Ascent of sap.

Composition of Xylem Sap

Xylem sap is very clear (nearly viscosity-1.0), slightly
acidic (pH-5.0). Most of the xylem sap is water and it
contains little organic molecules like amino acids,
organic acids, carbohydrates, inorganic molecules (ions
or NO3, SO4, PO4, Fe, Mg, Mn, etc.). the concentration of
the solute is even less than 1%. How ever it may vary
with the type of plant and environmental conditions.

Mechanism of Ascent of Sap

Several theories have put forwarded to explain the
mechanism of Ascent of Sap. Among which the
transpiration pull theory of Dixon and Jolly is the
acceptable one- as it is having maximum experimental
evidences compared to other theories.

Transpiration Pull Theory/ Cohesion Tension theory

Dixon and Jolly 1894 proposed Transpiration Pull Theory, Later
supported by Renner, Curtis, Clark and others.
The main features of this theory are as follows.

Cohesion (Attraction + Adhesion) of water molecules:

Water molecules have a strong cohesive force by which they are
held tightly to one another and cannot be easily separated.
Adhesion force of water molecules:
The water molecules have an adhesive property, which makes them
a strong attraction to the inner wall of xylem. Hence water
molecules cannot be easily torn away from the wall of the xylem.

Continuity of water column:

The entire water from root hair cells to the mesophyll cells of the
leaf is a continuous chain.
The water column present in different xylem vessels is not
independent as they are connected laterally.
Any influence in upper or lower parts is felt by entire column.

Transpiration pull or Suction force:

The water network as two terminals, one at the root tip
and other at the stomatal region of mesophyll of leaves.
The water evaporates from mesophyll cells due to
transpiration. This increase the DPD, which results water
flow from adjacent mesophyll cells.
Now the DPD of adjacent mesophyll cells increase, which
results water flow from inner mesophyll cells.
Similarly DPD of inner mesophyll cells increase, which
results water flow from xylem elements.
Continued transpiration in the leaf creates a tension in
the xylem elements in upper parts.
Since the water column is continuous and unbroken this
is transported down in to the root up to the area of
absorption and water moves up due to this force.

Merits of Transpiration Pull Theory

The theory as much experimental evidences when
compared to other theories.
It is very simple, logical, fundamental, easily
understood, acceptable and can be easily
demonstrated.

Demerits of Transpiration Pull Theory

The continuity of water column is in view of several
physical barriers in the form of tissues is doubtful.
Presence of air bubble in the xylem sap.
Ascent of sap is very negligible when there is no
transpiration.
Cohesion force with stands the opposing forces gravitational and transpiration tension.

Loss of Water in plants

Terrestrial plants absorb soil water from its

root system. Only little quantity (1-4%) is

used for the metabolic activities and most
(96-99%) is loosed in the form of water
vapors (Transpiration) or water droplets
(Guttation).
Transpiration
Transpiration may be defined as the process
of loss of water in the form of water vapors
by the aerial parts of the body in plants.

Types of transpiration:
Transpiration is of three types namely
Lenticular transpiration: the transpiration occurring
through the Lenticels is called as lenticular transpiration. It
accounts to 0.1% of total transpiration.
Lenticels-the openings formed in woody plants during secondary

growth to provide aeration to the inner living cells.

Cuticular transpiration: the transpiration occurring

through the cuticle is called cuticular transpiration. It

accounts to 3-10% of total transpiration.
Cuticle: the surface of the leaves and stems of certain plants are

covered with waxy coat to prevent he loss of water. This waxy coat
is regarded as cuticle. It is thick in regions of plants that are
exposed to sunlight and thin in the regions that are exposed to
shade.

Stomatal transpiration: the transpiration occurring

through the Stomata is called as stomatal transpiration. It

accounts to 90-95% of total transpiration.
Stomata: the small openings present in upper or lower or both the

surfaces of the leaves in the epidermal region.

Structure of Stomatal Apparatus

Stoma is a minute opening present in the epidermis of
leaf. The opening is usually surrounded by two
Crescent / Reniform cells called Guard cells.
The guard cells regulate the opening and closing of
stoma.
Guard cells cell wall is unevenly thick, usually thicker
towards stoma and thinner, elastic, Semi permeable
towards the outer region.
The cell wall is made from cellulose and pectin.
Each guard cell has a cytoplasmic lining, uninucleate
and contains chloroplasts.
The guard cells are surrounded by two or more
specialized cells called Subsidiary (Accessory) cells.
The stoma, guard cells and subsidiary cells together
constitutes the Stomatal complex (stomatal
apparatus).

Role of Stomata:
The guard cells regulate the opening and closing of
stoma, which in turn involved in Transpiration (90%).
It helps in the gaseous exchange during respiration
and photosynthesis.
It plays a key role in Passive absorption of water,
Ascent of Sap and absorption of mineral solutes.

Mechanism of stomatal movement

The guard cells open the stoma during the daytime
and closes during the night times, which is said to be
the stomatal movement.
The basic concept involved in opening and closing of
stoma is due to difference in osmotic potential that is
leading to turgidity or flaccidity.
Many theories are put

forwarded to explain
the mechanism of
stomatal movement.
The following are of
importance.

Starch Sugar Hypothesis:

Kohl (1895) proposed this theory. Later supported by Lloyd,
Searly and other scientists.
According to this theory,
During daytime photosynthesis decreases the production of CO2,
which in turn results in the increase in pH (neutral-7.0)
As a result the enzyme Starch phosphorylase gets activated and
convert starch in to Glucose- 1-Phosphate.
The formation of Glucose-1-Phosphate results in endosmosis,
further leads to turgid condition due to which the Stoma opens.
In the night the entire process is exactly reversed resulting in the
closure of Stoma.

Objections:
Reversal of Phosphorylase activity is doubtful.
The formation of Glucose-1-Phosphate may not cause mach
change in osmotic potential.
Correlation between the rate of Starch conversion and opening is
not established.

Stewards modified theory (1964)

To over come the objections Steward modified the above theory.
According to this
Glucose-1-Phosphate should have its phosphate removed to be
ultimately able to increase the concentration of Cell sap. The
process of opening and closing of stomata is steward theory is as
follows
Opening of Stomata

Closing of stomata

Objections:
Some guard cells do not have starch and they have some other
polysaccharides.
No evidence to show the presence of sugar at the time when starch
disappears when stomata is open.
In some starch is formed at a pH found during the opening of
stomata.

Potassium Ion pump theory:

Zelitch (1963) and Kevill and others proposed the potassium ion
pump theory. According to this theory the opening and closing
of stomata is due to influx and efflux of K+ and other ions. The
theory is summarized as followsOpening of Stomata
The potassium ions are pumped by the K+ pump by the

stimulation of Active cytokinin hormone

The CO2 concentration falls, pH raises and starch is broken

down to for malate, which restores the electrical equilibrium.

K+ and malate accumulates in the vacuoles in guard cells;

water flows in to guard cells; causing high turgor pressure in

these cells.
Guard cells expand asymmetrically and stoma opens.

Closing of stomata:
The movement of K+ stopped due to the phytohormone

Abscisic acid
The CO2 concentration raises, pH decreases and malate

reforms the starch.

Factors influencing the activity of Stomata:

Concentration of CO2: As the concentration increases the

stomata are closed.

Light: In high intensity and absence of light the stoma closes

and opens during moderate intensity light (Blue and Red light).
Water stress: when water is scarce the stomata is closed.
Temperature: At high and Low temperature the stoma closes and

min moderate temperature stoma opens.

Factors influencing the rate of transpiration

Humidity: As the humidity increases the rate of transpiration

decreases.
Temperature: relative temperature increases the rate of

transpiration. High or low temperature decreases the rate of

transpiration.
Wind: As the wind activity is raised the rate of transpiration

increases.
Light: relative light (Blue and Red light) increases the rate of

Advantages of transpiration
Responsible for absorption of water.
Main force in Ascent of sap.
Mineral uptake.
Removal of excess of water.
Translocation of solutes from one part to the other.
Opening and closing of Stomata indirectly helps in
Respiration and Photosynthesis.

Disadvantages:
Excess of transpiration lead to the wilting of
plants, which in turn lead to injury of plants even
death.
It affects photosynthesis process.

Antitranspirants:
Artificial device/substance to check/reduce
transpiration in plants is called Antitranspirants.
Antitranspirants are useful in the cultivation pf crops.
While applying the care is taken not to affect the
gaseous exchange that directly influence
photosynthesis and respiration.
Colorless plastics, silicon oils, low viscous waxes
impermeable to water but permeable to CO 2 and O2.
Application of Phenyl mercuric acetate leads the
partial closure of stomata.
Phytohormone Abscisic acid leads to closure of
stomata.
Increase in the CO2 also leads to the partial closure
of stomata.

Guttation
The process of physical loss of water in the
form of water droplets through Hydathodes
is called Guttation.
Guttation occurs during high rate of water
absorption and low rate of transpiration.
Guttation is observed during early morning
after a warm and humid night.
Guttation does not occur in all plants.
Water lost by Guttation is not pure as it
contains mineral solutes.

Structure of Hydathodes:
Hydathodes are the
special openings (Pores)
present in the exterior
surface of the leaf
epidermis. Internal to pore
is Epithem, which is made
from loosely arranged thin
walled parenchyma cells.
Epithem usually occupies
the tip of a vein.
Surrounding the epithem
is the mass of
chlorenchymatous tissue.

Translocation of solutes
Translocation may be defined as the process of transportation of
solute materials in the solution form from the region of their
synthesis to the region of storage or utilization.
The translocation of solute takes place in linear (both upward and
down ward) and radial directions.
The translocation occurs in the phloem.

Evidences to show the phloem as the site of translocation

Girdling or Bark ringing experiment:
A healthy plant is taken and its ring of bark including phloem is
selectively removed, the plant is kept in sunlight to carry out
photosynthesis.
The downward translocation food material stops and the food
material accumulate above the ring.
After 2-3 days the region above the ring swells due to
accumulation of food material.
This proves the involvement of phloem in the translocation of
food.

Tracer technique / Radiography method

The plant is allowed to photosynthesize in an atmosphere
in the presence of radioactive 14 CO2.
The plant synthesizes the radioactive labeled sugars and it
is transported in the plant.
Then the section is taken and subjected to
autoradiography.
Which shows that the substances are translocated through
the Sieve tubes of the phloem.

Composition of phloem sap

The composition of phloem sap may vary from one plant
to another. However mostly it contains sucrose, little
quantities of other carbohydrates, amino acids, amides,
phyto hormones and inorganic elements like calcium,
magnesium, potassium, lithium, nitrogen etc.
Sucrose is the major element that is translocated in the
plant body.

The mechanism of translocation of

organic solutes
Several theories / hypothesis was
made to explain the mechanism of
translocation of organic solutes. No
theories are complete and full-fledged.
How ever the popular with least
objections is Munchs mass flow
Hypothesis.

Munchs mass flow Hypothesis (Pressure flow hypothesis)

Ernst munch (1930) proposed a hypothesis to explain the
mechanism of translocation of organic solutes. Later this is
elaborated by crafts (1938).
According to this theory the food materials in plants flow
through the sieve tubes from the supply end (mesophyll) to the
consumption end (non green parts). i.e. the roots and storage
organs.
The formation of sugars in mesophyll (due to photosynthesis)
increases the osmotic potential, causing absorption of water
from xylem to dissolve sugars, which in turn increase the
pressure potential.
At the same time, in the cells of the root consumes the sugars
and storage organs stores the sugars in the form of insoluble
starch. Due to this turgor pressure in this region is lowered.
As a result the water with dissolved solutes move passively
from the supply end to the consumption end. Munch used two
osmometers A and B, which are made from semi permeable
membranes to explain the translocation of solutes.

 Osmometers A and B were connected by bent tube

C. Osmometer A was filled with concentrated sugar

solution and B with dilute solution and the tube C
with water. The whole apparatus was immersed in a
trough containing water. The water enters in to
osmometer A due to osmosis and its turgor pressure
is increased this increase in the turgor pressure
causes the mass flow of sugar solution to
osmometer B through tube C under the influence of
turgor pressure gradient. The water from B is forced
out because of the pressure created. The movement
of solutes from A to B continues till the solution in
both
the
osmometers
attains
the
same
concentration. This process can be prolonged if a
continuous supply of sugar solution in osmometer A
is maintained.
Munch compared osmometer A to supply end

Vein loading:
Vein loading is an active mechanism of transferring
sugars from the mesophyll cells in to the Sieve tube
ends of vein in the leaves. This mechanism involves
consumption of metabolic energy.
This process, maintains a high concentration of
sugars at the rate leaf end of the tube the sugars
are unloaded from sieve tube ends in to the
receiving ends of the plants.
Certain cells called transfer cells are involved in the
process of vein loading. These cells are special
parenchymatous cells with an expanded internal
surface, which helps in active transport and loading
of solutes in to the sieve tube elements of phloem.
These cells are found in the leaf parenchyma
adjoining the sieve tube elements of phloem in the
vascular bundle.