TRANSPIRATION

• Although large quantities of water are

absorbed by plant from the soil but
only a small amount of it is utilized.
The excess of water is lost from the
aerial parts of plants in the form of
water vapours. This is called as
transpiration.
• Transpiration is of three types
 1. Stomatal transpiration
• Most of the transpiration takes place
through stomata. Stomata are usually
confined in more numbers on the lower
sides of the leaves.
• In monocots. Eg. Grasses they are
equally distributed on both sides. While
in aquatic plants with floating leaves
they are present on the upper surface.
2. Cuticular transpiration
• Cuticle is impervious/ impermeable to
water, even though, some water may be
lost through it. It may contribute a
maximum of about 10% of the total
transpiration.
3. Lenticular transpiration
• Some water may be lost by woody stems
through lenticells which is called as
lenticular transpiration.
• Stomatal transpiration - This is the most dominant form of
transpiration being responsible for most of the water loss in
plants. It accounts for 90-95% of the water transpired from
leaves. As the name suggests, the process involves the
participation of the stomata (sing. stoma) or stomates,
microscopic pores in the epidermis of the leaves.
• Cuticular Transpiration – This type of transpiration is
responsible for the loss of water in plants via the cuticle. Water
vapor directly diffuses through the cuticle on leaves and
herbaceous stems and escapes to the atmosphere. The cuticle is a
waxy or resinous layer of cutin, a fatty substance, covering the
outside (epidermis) of leaves and other plant parts. Except for the
interruption by stomates and lenticels, the layer is continuous.
• 3. Lenticular Transpiration – This type of transpiration is the
loss of water from plants as vapor through the lenticels.
The lenticels are tiny openings that protrude from the barks in
woody stems and twigs as well as in other plant organs.
 Mechanism of stomatal transpiration

• The mechanism of stomatal transpiration which

takes place during the day time can be studied in
three steps.
1. Osmotic diffusion of water in the leaf from xylem
to intercellular space above the stomata through
the mesophyll cells.
2. Opening and closing of stomata (stomatal
movement)
3. Simple diffusion of water vapours from intercellular
spaces to other atmosphere through stomata.
• Inside the leaf the mesophyll cells are in
contact
• With xylem, and on the other hand with
intercellular space above the stomata.
• When mesophyll cells draw water from the
xylem they become turgid and their diffusion
pressure deficit (DPD) and osmotic pressure
(OP) decreases with the result that they release
water in the form of vapour in intercellular
spaces close to stomata by osmotic diffusion.
• Now in turn, the O.P and D.P.D of
mesophyll cells become higher and
hence, they draw water form xylem
by osmotic diffusion.
Opening and closing of stomata (Stomatal
movement)
• The stomata are easily recognized from
the surrounding epidermal cells by their
peculiar shape.
• The epidermal cells that immediately
surround the stomata may be similar to
other epidermal cells or may be different
and specialized. In the latter case, they are
called as subsidiary cells.
• The guard cells differ from other epidermal
cells also in containing chloroplasts and
peculiar thickening on their adjacent surface
(in closed stomata) or on surfaces.
• Consequent to an increase in the osmotic
pressure (OP) and diffusion pressure deficit
(DPD) of the guard cells (which is due to
accumulation of osmotically active
substances), osmotic diffusion of water from
surrounding epidermal cells and mesophyll
cells into guard cells follows.
• This increase the turgor pressure
(TP) of the guard cells and they
become turgid.
• The guard cells swell, increase in
length and their adjacent thickened
surfaces start forming a pore and
thus the stomata open.
• The thickened surfaces of the guard
cells come close to each other,
thereby closing the stomatal pore
and stomata.
• On the other hand, when OP and DPD
of guard cells decrease (due to
depletion of osmotically active
substances) relative to surrounding
epidermal and mesophyll cells, water is
released back into the latter by osmotic
diffusion and the guard cells become
flaccid.
• Osmotic diffusion of water into
guard cells occur when their
osmotic pressure increases and
water potential decreases (i.e
become more negative) related
to those of surrounding
epidermal and mesophyll cells.
• The guard cells become flaccid
when their osmotic pressure
decreases relative to the
surrounding cells (Movement of
water takes place from a region of
higher water potential to a region of
lower water potential.
These may be several different agents or mechanisms which
control stomatal movements.
1. Hydrolysis of starch into sugars in guard cells
2. Synthesis of sugars or organic acids in them
3. The active pumping of K+ ions in the guard.

• Hydrolysis (/haɪˈdrɒlɪsɪs/; from Ancient Greek

hydro-, meaning 'water', and lysis, meaning 'to
unbind') is a term used for both an electro-chemical
process and a biological one. The hydrolysis of
water is the separation of water molecules into
hydrogen and oxygen atoms (water splitting) using
electricity (electrolysis).
 1. Hydrolysis of starch into sugars in guard cells

Starch – sugar Inter conversion theory

• This classical theory is based on the
effect of pH on starch phosphorylase
enzyme which reversibly catalyses
the conversion of starch + inorganic
phosphate into glucose -1
phosphate.
• During the day, pH in guard cells is
high. This favours hydrolysis of
starch (which is insoluble) into
glucose -1- phosphate (which is
soluble) so that osmotic pressure is
increased in guard cells.
• Consequently water enters into the guard
cells by osmotic diffusion from the
surrounding epidermal and mesophyll cells.
Guard cells become turgid and the stomata
open.
• During dark, reverse process occurs. Glucose
1- phosphate is converted back into starch in
the guard cells thereby decreasing osmotic
pressure. The guard cell release water,
become flaccid and stomata become closed.
According to Steward (1964), the
conversion of starch and inorganic
phosphate into glucose-1-phosphate does
not cause any appreciable change in the
osmotic pressure because the inorganic
phosphate and glucose-1-phosphate are
equally active osmotically.
In this scheme he has suggested that,
• Glucose-1-phosphate should be further
converted into glucose and inorganic
phosphate for the opening of stomata.
• Metabolic energy in the form of ATP
would be required for the closing of
stomata which probably comes through
respiration.
 2. Synthesis of sugars or organic acids in
Guard cells
• During day light photosynthesis occurs
in guard cells as they contain
chloroplast. The soluble sugars formed
in this process may contribute in
increasing the osmotic potential of
guard cells and hence resulting in
stomatal opening.
• However, very small amounts of soluble
sugars (osmotically active) have been
extracted from the guard cells which are
insufficient to affect water potential.
• As a result of photosynthesis CO2
concentration in guard cells decreases
which leads to increased pH up of organic
acids, chiefly malic acid during this period
in guard cells.
• The formation of malic acid would
produce proton that could operate in an
ATP-driven proton K+ exchange pump
moving protons into the adjacent
epidermal cells and K ions into guard cells
and thus may contribute in increasing the
osmotic pressure of the guard cells and
leading to stomatal opening.
• Reverse process would occur in darkness.
 3. ATP –Driven proton (H+) – K exchange
pump mechanism in Guard cells
• According to this mechanism, there is
accumulation of K+ ions in the guard
cells during day light period.
• The protons (H+) are ‘pumped out’ from
the guard cells into the adjacent
epidermal cells and in exchange K+ ions
are mediated through ATP and thus are
an active process.
• ATP is generated in non-cyclic
photophosphorylation in
photosynthesis in the guard cells. The
ATP required in ion exchange process
may also come through respiration.
• The accumulation of K+ ion is sufficient
enough to significantly decrease the
water potential of guard cells during day
light.
• Consequently, water enters into them
from the adjacent epidermal and
mesophyll cells thereby increasing
their turgor pressure and opening the
stomatal pore.
• Reverse situation prevails during dark
when stomata are closed. There is no
accumulation of ‘K’ in guard cells in
dark.
• (iii) The last step in the mechanism of
transpiration is the simple diffusion of
water vapours from the intercellular
spaces to the atmosphere through open
stomata.
• This is because the intercellular spaces
are more saturated with moisture in
comparison to the outer atmosphere in
the vicinity of stomata.
 Significance of Transpiration
• Plants waste much of their energy in
absorbing large quantities of water and
most of which is ultimately lost through
transpiration.
• Some people think that – Transpiration
as advantageous to plant. Others regard
it as an unavoidable process which is
rather harmful.
 Advances of transpiration
1. Role of movement of water
• Plays an important role in upward movement
of water i.e Ascent of sap in plants.
2. Role in absorption and translocation of
mineral salts
• Absorption of water and mineral salts are
entirely independent process. Therefore
transpiration has nothing to do with the
absorption of mineral salts.
• However, once mineral salts have
been absorbed by the plants, their
further translocation and
distribution may be facilitated by
transpiration through
translocation of water in the
xylem elements.
• which raises their temperature.
Transpiration plays an important role in
controlling the temperature of the
plants.
• Rapid evaporation of water from the
aerial parts of the plant through
transpiration brings down their
temperature and thus prevents them
from excessive heating.
 Transpiration as a necessary evil
1. When the rate of transpiration is high
and soil is deficient in water, an internal
water deficit is created in the plants
which may affect metabolic processes
2. Many xerophytes have to develop
structural modification and adaptation
to check transpiration.
3. Deciduous trees has to shed their
leaves during autumn to check loss of
water.
• But, in spite of the various
disadvantages, the plants cannot
avoid transpiration due to their
peculiar internal structure particularly
those of leaves.
• Their internal structure although
basically mean for gaseous exchange
for respiration, P.S. etc. is such that it
cannot check the evaporation of
water.
• Therefore, many workers like Curtis
(1926) have called transpiration as
necessary evil.
 Factors affecting transpiration rate

A. External factors
1. Atmospheric humidity
• In humid atmosphere, (when relative humidity
is high), the rate of transpiration decreases. It
is because atmosphere is more saturated with
moisture and retards the diffusion of water
vapour from the intercellular spaces of the
leaves to the outer atmosphere through
stomata.
• In dry atmosphere, the RH is low and the
air is not saturated with moisture and
hence, the rate of transpiration increases.
2. Temperature
• An increase in temperature brings about
an increase in the rate of transpiration by
1. lowering the relative humidity
2. Opening of stomata widely
3. Wind
i. When wind is stagnant (not blowing), the
rate of transpiration remains normal
ii. When the wind is blowing gently, the rate of
transpiration increases because it removes
moisture from the vicinity of the
transpiration parts of the plant thus
facilitating the diffusion of water vapour
from the intercellular spaces of the leaves to
the outer atmosphere though stomata.
• iii. When the wind is blowing
violently, the rate of transpiration
decreased because it creates
hindrance in the outward diffusion
of water vapours from the
transpiring part and it may also
close the stomata.
4. Light
• Light increases the rate of transpiration
because, In light stomata open; It increases
the temperature. In dark, due to closure of
stomata, the stomatal transpiration is almost
stopped.
5. Available soil water
• Rate of transpiration will decrease if there is
not enough water in the soil in such from
which can be easily absorbed by the roots.
6. CO2
• An increase in CO2 concentration in
the atmosphere (Over the usual
concentration) more so inside the
leaf, leads towards stomatal closure
and hence it retards transpiration.
 B. Internal factors
1. Internal water conditions
• It is very essential for transpiration.
Deficiency of water in the plants will result
in decrease of transpiration rate.
• Increase rate of transpiration containing
for longer periods often create internal
water deficit in plants because absorption
of water does not keep pace with it.
2. Structural features
• The number, size, position and the movement
of stomata affect rate of transpiration.
• In dark stomata are closed and stomatal
transpiration is checked. Sunken stomata help
in reducing the rate of stomatal transpiration.
• In xerophytes the leaves are reduced in size or
may even fall to check transpiration. Thick
cuticle on presence of wax coating on exposed
parts reduces cuticles transpiration.
 Anti-transpirants
• A number of substances are known which
when applied to the plants retard their
transpiration. Such substances are called
as anti-transpirants.
• Some examples of anti-transpirants are
colourless plastics, silicone, oils, low
viscosity waxes, phenyl mercuric acetate,
abscisic acid, CO2, etc.
• Colourless plastic, silicone oils and low
viscosity waxes belong to one group as these
are sprayed on the leaves, form after film
which is permeable to O2 and CO2 but not to
water.
• Fungicide phenyl mercuric acetate, when
applied in low concentration (10-4 m), it
exercised a very little toxic effect on leaves
and resulted in partial closure of stomatal
pores for a period of two weeks.
• Similarly Abscisic acid (ABA) a plant
hormone also induces stomatal closure.
• CO2 is an effective antitranspirants. A little
rise in CO2 concentration from the natural
0.03% to 0.05% induces partial closure of
stomata. Its higher concentration cannot
be used which results in complete closure
of stomata affecting adversely the
photosynthesis and respiration.
 GUTTATION
• In some plants such as garden nasturtium,
tomato, colocasia etc, water drops ooze
out from the uninjured margins of the
leaves where a main vein ends.
• This is called as guttation and takes place
usually early in the morning when the rate
of absorption and root pressure are high
while the transpiration is very low.
• The phenomenon of guttation is
associated with the presence of special
types of stomata at the margins of the
leaves which are called as water stomata
or hydathodes.
• Each hydathode consists of a water pore
which remains permanently open. Below
this there is a small cavity followed by a
loose tissue called as epithem.
• This epithem is in close association with
the ends of the vascular elements of veins.
Under high root pressure the water is
given to the epithem by the xylem of the
veins.
• From epithem water is released into the
cavity. When this cavity is completely filled
with watery solution, the later begins to
ooze out in the form of watery drops
through the water pore.