Plant Water Relations

By Dr Kabir Pokhrel C/B online class 2022/23
PLANT WATER RELATIONS
@ Physiology……………………………………………
@ Father of plant physiology……………………………………………
@ Father of Indian plant physiology: Sir J.C. Bose

@ Water is very essential factor for carrying out all the physiological
functioning of plants.
@ Transport is the process of moving water, minerals and food to all parts
of the plant body.
@ Conducting tissues such as xylem and phloem play an important role in this.
@ Based on the distance travelled by water (sap) or food (solute) they are
classified as
1. Short distance (Cell to cell transport) and

2. Long distance transport.
@ Short-distance (Cell to cell transport):

……………………………………….
o Examples: Diffusion, Imbibition, and Osmosis.
@ Long-distance transport:
……………………………………………………………
o Examples: Ascent of Sap and Translocation of Solutes.
@ Based on energy expenditure during transport, transport may be passive
transport and active transport.
@ Active transport: It is a biological process and it runs based on the energy
obtained from respiration. It is an uphill process.
@ Passive transport: It is a downhill process which utilizes physical forces
like gravity and concentration. No energy expenditure is required. It
includes diffusion, facilitated diffusion, imbibition, and osmosis.
Diffusion:
@ The net movement of molecules from a region of their higher concentration
to a region of their lower concentration along a concentration gradient
until an equilibrium is attained.
@ Diffusion is a passive process, which takes place along the concentration
gradient of solution. [BPKIHS]
@ Diffusion occurs due to the kinetic energy of particles of a matter.
@ Rate of diffusion:
• ………………………….
• ……………………………
• ………………………………
Characteristics of diffusion
• It is a passive process, hence no energy expenditure involved.
• It is independent of the living system.
• Diffusion is obvious in gases and liquids.
• Diffusion is rapid over a shorter distance but extremely slow over a longer
distance.
• The rate of diffusion is determined by temperature, concentration gradient
and relative density.

Significance of diffusion in Plants
• Gaseous exchange of O2 and CO2 between the atmosphere and stomata of
leaves takes place by the process of diffusion. O2 is absorbed during
respiration and CO2 is absorbed during photosynthesis.
• In transpiration, water vapour from intercellular spaces diffuses into

atmosphere through stomata by the process of diffusion.
• The transport of ions in mineral salts during passive absorption also takes
place by this process.
@ When diffusion of two or more substances takes place simultaneously at
the same time, it is called ……………………………….. [NEET 2016]
Faciliated Diffusion:
@ In facilitated diffusion, molecules cross the cell membrane with the help
of special membrane proteins called transport proteins, without the
expenditure of ATP.
@ Two types of transport proteins present in the cell membrane are channel
protein and a carrier protein.
@ Channel Protein : Channel protein forms a channel or tunnel in the cell

membrane for the easy passage of molecules to enter the cell. Examples:
Porin and Aquaporin.
• Porin: large transporter protein found in the outer membrane of
plastids, mitochondria and bacteria
• Aquaporin: water channel protein embedded in the plasma membrane.
Example: proximal tubule and descending thin limb of nephron
@ Carrier Protein: Carrier protein acts as a vehicle to carry molecules
from outside of the membrane to inside the cell and vice versa.
1. Uniport: molecule of a single type move across a membrane

independent of other molecules in one direction.
2. Symport or co-transport: transports two types of molecules
across the membrane in the same direction.
3. Antiport or Counter Transport: simultaneously transports two
different molecules, in opposite directions, across the membrane.
Diffusion pressure and diffusion pressure deficit (DPD)

@ Diffusion pressure is hypothetical term which describes the potential
ability of a gas, liquid or solid to diffuse from its higher concentration to
lower concentration.
@ Every liquid has a definite diffusion pressure. A pure solvent has
……………. diffusion pressure. [BPKIHS, IOM]
@ When some amount of solute is added to it, its diffusion pressure
…………….
@ This deficit in diffusion pressure of the solution due to the addition of
solute is called ………………..
Or
@ DPD………………………………………………………………..
@ Diffusion pressure deficit (DPD) is the index of absorbing power of a

solution. DPD is the main force responsible for the absorption of water by
plants. It is also called …………………………………..
@ A cell absorbs water till its DPD becomes zero.
DPD µ Concentration of the solution

Question: Maximum value of DPD can be observed in:
a. Leaf cell b. Stem cell
c. Root cell d. Soil water
Hint: Remember that water moves from a solution of low DPD to the solution
of high DPD. We know that the flow of water is from:
Soil Root cell Stem cell Leaf cell
So the order of increasing DPD will be:

Soil water < Root cell < Stem cell < Mesophyll (leaf) cell
@ Also remember that water moves from a solution of high water potential
to the solution of low water potential.
So order of increasing water potential will be:
Mesophyll cell < Stem cell < Root cell < Soil water
Types of Solution
1. Hypotonic solution: ………………………………………………….
2. Hypertonic solution: ………………………………………………..

3. Isotonic solution: …………………………………………………
Figure: Effect of different types of solution on plant and animal cell

Types of membrane
Figure: Types of membrane & movement of substance across different

membrane
1. Permeable membranes: Such membranes or walls allow diffusion of both,

solvent and solute molecules or ions through them, e.g. cellulose wall of
cells, lignified cell walls etc.
2. Impermeable membranes: Such membranes prohibit the diffusion of both,
solvent and solute particles through them, e.g. heavily cutinised or
suberised cell walls in plants.
3. Semipermeable membranes: Such membranes allow diffusion of solvent
molecules but do not allow the passage of solute molecules, e.g. membranes
of collodion, parchment paper and copper ferrocyanide membranes.
[BPKIHS]
4. Differentially permeable membranes: These membranes allow some
selective solutes to pass through them along with the solvent molecules.
Such membranes are also called selectively permeable membranes, e.g.
……………………………………………………………………..
Osmosis
@ Osmosis is special type of diffusion which involves the movement of
……………… or ……………………… from the region of their
higher concentration to a region of their lower concentration through a
………………………………… membrane.
@ Note:
• Movement of minerals through a cell membrane is also diffusion (not
osmosis).
• Movement of water molecules from higher concentration to lower
concentration without semipermeable membrane is also diffusion.
Figure: Demonstration of Osmosis

@ In osmosis water moves from:
• …………………………………………….
• ………………………………………………..
• …………………………………………………
• ………………………………………………….
@ Osmosis continues until hydrostatic pressure becomes equal to osmotic
pressure.
@ Exosmosis is the movement of solvent from cell to surrounding solution
(hypertonic). Exosmosis results ……………………………
@ Endosmosis is the movement of solvent from surrounding solution
(Hypotonic) to cell. Endosmosis results in ……………………………..
Osmotic pressure (O.P.)

@ Pressure required to stop osmosis when a solution is separated from its
pure solvent by a semi-permeable membrane is known as osmotic pressure.
• O.P. µ ……………………………………….
• O.P. of pure water is ………………………
• Hence O.P. is always positive for a solution.
@ Highest O. P. is found in halophytes and minimum O.P. is found in

hydrophytes. Increasing order of O. P. in plants:
………………………………………………………….
@ Maximum osmotic pressure has been recorded in ……………………….
@ The osmotic pressure of a solution is measured by ……………….
@ O.P. of cell is measured by incipient plasmolysis.

@ O.P. of electrolyte is more than O.P. of non-electrolytes. For example:
Solution of 1 M NaCl is higher than O.P. of 1 M glucose.
A B A B
(1) 2M 2M (2) 4M 2M
NaCl Sucrose NaCl AlCl3
A ¬ B
No movement because A and B has equal no. of
because, 1M Nacl Na + Cl¯ i.e. 2 moles of ions
+
ions.
but 1M sucrose = 1M sucrose because it does
not dissociate
Concept of Water Potential:

@ It is defined as amount of free energy present per mole of water
molecule.
@ Water potential is represented by Greek letter 'y' (Psi) and expressed
in bars.
@ Water potential (y) of pure water is …………………. and is assumed
to be zero and addition of solute in it, decreases its water potential (i.e.
negative value). [NEET 2017]
@ Water potential (y) of protoplasm is equal but opposite in sign to D.P.D.
i.e., Water potential = -D.P.D.

Turgor pressure (T.P.) and Wall pressure (W.P.)

@ Turgor pressure (T.P.) is the pressure exerted by ……………………
• Turgor pressure is a hydrostatic pressure with which cell sap presses

the cytoplasm against cell wall.
• Turgor pressure develops only inside the osmotic system and is not
applicable for free solution.
• Turgor pressure acts against osmotic entry of water into cell while
osmotic pressure helps in the osmotic entry of water into the cell.
• A flaccid cell has zero turgor pressure.
@ Wall pressure (W.P.) is pressure exerted by …………………………
………………………………………….. due expanding protoplasm.

• Wall pressure is always equal and opposite of ………………………
@ Plant cell does not burst, when placed in pure water due to …………….,
@ An animal cell burst when placed in pure water because wall pressure is
absent.
Figure: Cell showing T.P., W.P. and O.P.

• In fully turgid cell: Osmotic pressure = Turgor pressure
[IOM/MOE/BPKIHS/IE]
@ Protoplasm of the cell attached with the cell wall is due to turgidity of the
cell.
@ Plant movement like, movement of guard cells of stomata, wilting movements
and seismonastic movements etc. depend upon turgor pressure.
@ Turgor pressure keeps the leaves, floral parts etc. in stretched condition.
Osmotic relationship between DPD, T.P. and O.P.:
@ The diffusion pressure of a pure solvent is always more than that of its
…………………..
@ The amount by which the diffusion pressure of a solution is lower than
that of its pure solvent is called …………………………………….
@ DPD determines the direction of osmosis and it is the power of absorption
of water for the cell (Suction pressure)
@ Normally, osmotic pressure is greater than the turgor pressure in a cell.
The difference between osmotic pressure and turgor pressure is called
suction pressure or DPD.
DPD = OP – TP (WP)
DPD for fully turgid cell:

@ When a cell is placed in pure water or hypotonic solution then water enter
into the cell, results turgor pressure develop in the cell. The cell starts
swelling due to the turgor pressure. Simultaneously, concentration of cell
sap decreases due to continuous inflow of water.
@ Therefore OP goes on decreasing and T.P. goes increasing, when value of
TP will be equal to the OP then DPD will be zero.
@ At this stage cell becomes …………………………...
Therefore in a fully turgid cell:

DPD = …………………………………………………………
…………………………………………………………………
DPD in flaccid cell:
@ If, the cell is in flaccid state then its T.P or WP would be zero and value
of DPD would be equal to OP.
TP or WP = O
DPD or S.P. = OP
Therefore, if a flaccid cell placed in water then waters enter into cell
because DPD of the cell sap is higher.
DPD for plasmolysed cell:
@ Sometimes the value of turgor pressure is negative as in plasmolysed cell.
In this state DPD = OP – TP
DPD = OP – [- TP] = OP + TP
Therefore, DPD of the plasmolysed cell is greater than osmotic pressure.
Demand of water= Plasmolysed cell> Flacid cell> Partially turgid cell>
Fully turgid cell
@ D.P.D can never be negative.
@ D.P.D. is a Driving force of the absorption in water in roots.
@ In normal condition OP > TP
Question:
If cell A with OP = 5 and TP = 4 is surrounded by cells with OP = 3 and
TP = 1. What will be the direction of water movement?
a. From cell A to other cells b. From other cells to cell A
c. Water will not move d. Water will move up
Osmotic pressure in a vacuolated plant cell is:
a. Equal to W.P. b. Equal to T.P.
c. More than D.P.D. d. Less than D.P.D.
Exosmosis and Plasmolysis:
@ Plasmolysis is the shrinking of protoplast due to exosmosis when cell is
placed in hypertonic solution. Plasmolysis is the result of exosmosis. [IOM
1998]
@ The point at which plasmolysis first appears is called incipient point and
plasmolysis at this point is incipient plasmolysis.
@ The space between the cell wall and shrinked cytoplasm is occupied by
outer hypertonic solution.[IOM, BPKIHS]
@ In incipient plasmolysis, cell membrane just separates from cell wall.
Significances of Plasmolysis
@ Plasmolysis is the best and quickest physiological process to identify
weather a cell is living or dead. Plasmolysis is not possible in dead cell.
@ Permeable nature of cell wall and semi-permeable nature of cell
membrane can be determined by plasmolysis. During plasmolysis, there is
no shrinkage of cell wall because cell wall is rigid and gets stretched
@ Excess supply of fertilizers kills crops due to plasmolysis. [IOM 1996]
@ Bacteria cannot survive in salted pickles due to plasmolysis

@ Salt spread in tennis court kills weeds by plasmolysis.
Cell placed in
pure water
(Hypotonic
solution)
Result:
Increase turgor
pressure
Cell placed in
strong salt
solution
(Hypertonic
solution)
Normal cell on isotonic solution
Result:
Plasmolysis

Figure: Plasmolysis
@ When a cell is plasmolysed after being kept in a hypertonic solution, a

space is created between the cell wall and shrunken protoplasm. This
space contains ……………………………………... [I. E. 2009]
@ A plasmolysed cell regains normal condition, if placed in …………..
solution, it is called ……………………..
Imbibition
@ Imbibition is a physical process by which solvent is adsorbed by certain
substances making them swell. The solvent, which is adsorbed, is called
………………………..
@ The solute which adsorbs is called ……………………. Imbibition is a
phenomenon of ………………………... [BPKIHS 2006]

@ Imbibition process involves movement of water into imbibant through
diffusion as well as capillary action.
@ Imbibition is also diffusion, since water movement is along a concentration
gradient.
@ Only the solid substances containing …………………… can absorb
water. Swelling of dry seeds when placed in water is due to imbibition.
@ During imbibitions:
• ……………………………………
• …………………………………….
• ………………………………………
@ Heat evolved during imbibitions process is called ‘heat of hydration’ or
‘heat of wetting’.
@ Seed germination is accompanied by evolution …………………….
@ Agar-agar is a very efficient imbibant. It imbibes 99 times its mass of water

and is therefore used in bacterial culture.
@ Agar-agar is a phyco-colloid obtained from red algae: Gallidium,

Gracilaria and Gigartina (Agarophytes) [I. E. 1998]
@ Decreasing order of imbibition capacity:
…………………………………………………………………….
@ Rubber never imbibes water it only imbibes ether and other organic
solvents like kerosene. [BPKIHS 2004]
Significance of imbibition:
@ First physiological process during seed germination imbibition.
@ Initial process of water absorption in root hairs by root hairs is imbibition.
@ The pressure that is produced by swelling of wood had been used by
prehistoric man to split rocks and boulders.
@ During rainy season, wooden doors swell due to imbibition.
Absorption of Water by Plants:

Soil Texture
@ The property of soil based on the size of its particles is called texture.
@ Study of soil is known as “Pedology or Edaphology”.
@ Soil can be classified into four main types on the basis of the size of soil
particles:
Type Size of particle
Thus the correct sequence
Clay Less than 0.002 mm according to increasing particle
size is:
Silt 0.002-0.02 mm Clay < Silt < Fine sand < Coarse
sand < Gravel
Fine sand 0.02-0.2 mm
Coarse sand 0.2-2 mm
@ The soil transported by wind is referred to as Eolian soil and those

transported by action of gravitational force is called Colluvial deposits.
@ On the basis of proportion of different sized particles, soils are classified
into:
• Sandy soil (85% sand + 15% clay or silt or both)

• Loamy soil (50% sand + 50% clay or silt or both)
• Silt soil (90% silt + 10% sand)
@ “A horizon” is the topmost soil having maximum humus.
@ Clayey soils have poor aeration but maximum water holding capacity.
@ The least porous soil is clay soil. [BPKIHS 1997]
@ Loamy soil is considered best for cultivation of crop as it maintains an

equilibrium between the soil air and soil water into pore spaces and
contains all types of macro and micro elements required by plants for
their metabolism and growth. [I. E. 1999]
Pathway of Water Movement in Plants

@ Mainly plants absorb water through root hairs. Root hairs are present in
the zone of cell maturation.
Note: Maximum mineral absorption takes place through the zone of cell
elongation.
@ Root hairs are unicellular structures that arise from piliferous cells of
epiblema exogenously.
Soil solution Root hairs Epiblema Cortex
Metaxylem Protoxylem Pericycle cells Endodermis
@ Pathway followed by water is of two types, i.e., symplast pathway and

apoplast pathway.
• In the apoplast pathway, water moves exclusively through the cell wall
without crossing any membranes. The apoplast is the continuous system

of cell walls and intercellular air spaces in plant tissues.
• In the symplast pathway, water travels from one cell to the next via
the plasmodesmata. The symplast consists of an entire network of cell

cytoplasm interconnected by plasmodesmata.
Apoplastic
pathway
Symplastic
pathway
Epiblema Cortex Endodermis Xylem
Figure: Pathway of water movement

@ At the endodermis, water movement through the apoplast pathway is
obstructed by the Casparian strip.
@ Casparian strip is a band of radial cell walls in the endodermis that is
impregnated with the wax like, hydrophobic substance called suberin.
Suberin acts as a barrier to water and solute movement.
@ Casparian strip breaks the continuity of the apoplast pathway and forces
water and solutes to cross the endodermis by passing through the plasma
membrane.
@ Thus despite the importance of the apoplast pathway in the root cortex
and stele, water movement across the endodermis occurs through the
simplest.
Mechanism of water absorption:

@ Water is absorbed by two ways:
A. Passive absorption (water is absorbed through the root)
B. Active absorption (water is absorbed by the root)
Passive absorption of water:
@ According to this theory, the forces responsible for absorption of water
originate not in the cell of root but in the cells of transpiring shoots.
@ Due to transpiration, a tension is created in water in the xylem of the
leaves.
@ This tension is transmitted to the xylem of the roots through the xylem of
stem and water rises upward to the transpiring surface.
@ This type of absorption is known as passive absorption because the entry
of water into the root is brought about by the conditions, which originate in
the top of the plant (i.e, transpiration) and the root cells play only a
subsidiary role.
Active absorption of water:
@ Active absorption occurs as a result of activities in the root and does not
concern the shoot.
@ The force necessary for absorption of water is generated in the cells of
root itself (= absorption is by root).
@ This theory is supported by phenomena of root pressure and guttation. It
hold good in case of well-watered and slowly transpiring plant.
@ Two theories:
a. Active absorption through osmotic mechanism:

• According to this theory, root functions as an osmometer. In this type of
absorption, the osmotic pressure of cell sap of root hair is higher than
that of soil water.
• Therefore, water moves through the plasma membrane by osmotic
diffusion.
b. Active absorption through non-osmotic mechanism:

• Sometimes, absorption of water takes place even when the OP of soil
water is higher than that of cell sap.
• This type of absorption is against the osmotic gradient and known as
active non-osmotic absorption. The intake of water according to this
theory presumably requires the expenditure of energy released by
respiration. (BPKIHS 2012)
Differences between Active and Passive absorption

Active absorption Passive absorption
@ It accounts for 2 – 4 % of total @ It accounts for 96 – 98% of total
water absorption. water absorption.
@ It is at the expanses of ATP. @ No ATP needed.
@ It occurs in slowly transpiring @ It occurs in rapidly transpiring
plants. plants.
@ Rate of water absorption is low. @ Rate of water absorption is high.
@ Root pressure is developed. @ Shoot pressure is developed.
@ Root plays an active role. @ Shoot plays a passive role.
@ It involves symplast movement @ It involves both apoplast (through
(through protoplasm + cell wall + Intercellular space)
plasmodesmata) of water. and symplast movement of water.
@ Bleeding and guttation are @ Transpiration is associated with
closely associated with it. passive absorption.
Factors affecting water absorption by plants:

A. Available water content of the soil: Capillary water is the only water
which can be absorbed by plant roots [IE].
B. Soil temperature: Most plants require temperature ranging from 20o to 35o
for optimum water absorption.
Note:
• Low temperature soil is called psychrophilic soil, where physiological
dryness occurs. Due to this reason, plants growing on high altitudes are
xerophytic in habitat.
C. Concentration of soil solution: If the concentration of the soil solution is
more as compared to that of the cell sap of root, water absorption does not
take place, as the soil becomes physiologically dry.
Notes:
• Halophytes are adapted live and grow in soils where there is a high
concentration of salts. Such plants cells have high osmotic concentration

due to which their growth and survival becomes possible.
D. Soil aeration: Absorption of water is low in poorly aerated soil.
REMEMBER:
@ Rate of water absorption is:
• Decreased in saline soil (physiologically dry soil)
• Decreased in poorly aerated soil.
• Decreased in water logged soil.
• Is directly proportional to transpiration.
• Water logged soil shows physiological dryness due to anaerobic condition.
• Prolong water-logging kills plants due to stoppage of root respiration.
• Auxin treated cells can absorb water from a hypertonic solution by non-
osmotic absorption.
Ascent of sap
@ The upward movement of H2O with dissolved minerals through lumen of
xylem vessels against force of gravity is called ascent of sap.
@ Path of ascent of sap in gymnosperm is tracheids and in angiosperms is
mainly vessels.
@ That xylem is the principal water conducting tissue has been recognized at
least since the time of girdling experiments of Hartwig.
@ Best experiment to show path of ascent of sap is Ringing or Girdling

experiment.
@ In this experiment, a ring or girdle is made on stem by removing bark (all
the tissues outer to vascular cambium).
@ In this experiment; all tissue

outside vascular cambium is
removed in a ring.
@ Under this condition upper part
of phloem is connected to
lower part by only central
xylem cylinder and pith.
@ It results in accumulation
just above ring (=swelling)
and root dies first (=due to
Figure: Ringing (=girdling experiment) starvation).
v Ringing or girdling experiment
Theories of ascent of sap: cannot be performed in
monocots due to scattered
vascular bundles in stem.
Vital force theories: v
@ According to this theories, the forces required for ascent of sap is

generated in living cells.
@ These theories are not supported by experimental evidence hence they have
been discarded.
1. Relay pump theory given by Goldewiski is the first vital force theory
regarding ascent of sap. According to this theory, ascent of sap takes place
through xylem parenchyma.
2. Pulsation theory was introduced by sir J.C. Bose according to which
ascent of sap takes due to the pulsatory activity of living cells of
innermost cortical layer just outside the endodermis.
Root pressure theory:
@ Given by Priestley. Root pressure refers to positive hydrostatic pressure,
which sometimes develops in the xylem sap of roots as a result of metabolic
activities of roots.
Physical force theories:

1. Capillary force theory: By Boehm:
• According to this vessels & tracheids acts as capillaries & ascent of sap
takes place due to capillary force.
• Rise of water in the capillary continue till the force of surface tension is
counterbalanced by downward pull of gravity.
2. Imbibition force theory: By Unger & Von Sachs [MOE 2000]
• According to it, ascent of sap is due to the imbibition force of xylem wall.
3. Transpiration pull & cohesion force theory: By Dixon & Jolly
• Most accepted or universally accepted theory of ascent of sap.

• According to it 3 components are involved in ascent of sap.
• Cohesion: Mutual attraction between the water molecules is known as
cohesion, which form a continuous water column in xylem elements.
• Adhesion: Attraction between xylem walls & water molecules is called
adhesion force, which helps in maintenance of water column of xylem.
• Transpiration Pull: A tension or negative pressure develops in xylem,
due to rapid transpiration in leaves (because of high DPD), this creates
a transpiration pull, which is responsible for the pulling up of water
column in xylem. So ascent of sap is constitutive effect of cohesion,
adhesion & transpiration pull.
Water loss due to

transpiration
Leaf
Stem Xylem
Toward
root From roots
Ascent of sap Transpiration pull

(Water moving toward (Moving toward root)
leaves)
Figure: Transpiration pull (tension) and ascent of sap
Transpiration
@ Only small fraction of absorbed water is used by plant and remaining
(=nearly 98-99%) is lost in environment in vapour or liquid form.
@ Loss of water in the form water vapour from aerial parts of living plant
body is known as …………………………...
@ Whereas, the loss of water from aerial parts of living plants in the form
of liquid is known as …………………….
@ Driving force of transpiration is differences in vapour pressure between
the internal leaf spaces and outer atmosphere.
@ Transpiration is minimum in succulent xerophytes, maximum in mesophytes
@ "Transpiration
and absent in submerged evil" by Curtis.
hydrophytes.
is a necessary
@ “Transpiration is an unavoidable event” by Steward. [IOM 2000]
@ In herbaceous plant, 99% of total water absorbed by plants is lost through
transpiration. So, only 1 - 2% is actually used by them for their vital
functions.
Kinds of Transpiration
@ Transpiration is of following types.
1. Stomatal Transpiration
• ……………………………………………………….
• ……………………………………………………….
• ……………………………………………………….
• ……………………………………………………….
2. Cuticular Transpiration
• ……………………………………………………….
• ……………………………………………………….
• ……………………………………………………….
3. Lenticular Transpiration
• It occurs through the lenticles. About 0.1 to 1% of total transpiration is
lenticular. [NEET 2013]
Lenticular transpiration is found only in woody branches of the trees,
•
where leneticles are present.

4. Bark Transpiration
• It occurs through corky covering of the stems.
Stomatal Apparatus
@ The term stomata (Greek; Stoma = mouth) refers to a pair of guard cells
and pore between them.
@ Stomata are present not only in the epidermis of leaves but also in certain
fruits (e.g. banana, citrus, cucumber etc.).
@ Each stoma is surrounded by two ……………………………………
• Guard cells are rapidly influenced by turgor changes. [NEET 2017

• The concave-convex curvature of the two guard cells is variable and
causes the stomatal pore to open and close.
• The walls of guard cells are thickened on the inner side and thinner and
more elastic on the outer side. [IE 2006]
• Adjacent to each guard cells are accessory or subsidiary cells. [NEET

2016]
Figure: Stomatal apparatus; A. Aperture closed B. Aperture open

Quik Digest
• Guard cells are specialized epidermal cell containing chloroplast. These
have both photosystem I and II, but cannot assimilate CO2 because of
absence of Ribulose biphospate carboxylase (RUBISCO) and other enzymes.
• In dicot guard cells are reniform (kidney shaped) while monocot have dumb
bell shaped guard cells. @ DR MD [NEET 2018]
Distribution of Stomata
1. Hypostomatic (Apple or Mulberry type): Stomata are present only on lower
surface of leaf, e.g. Apple.
2. Epistomatic (Water lily type): Stomata are present only on the upper
epidermis of leaf, e.g., Water lily and many floating plants (Nelumbo,
Nymphaea, Victoria).
Note: In these plants, metabolism will be hindered if the leaves are
coated with wax on their upper surface. [I. E. 2001]
3. Amphistomatic: Stomata are present on both surfaces of leaves.
a. Anisostomatic (Potato type): Stomata are more on the lower surface. E.g.,
dorsiventral leaves
b. Isostomatic (Oat type): Stomata are equally distributed on both the
surfaces of leaf e.g., isobilateral leaves
5. Astomatic (Potamogeton type): Stomata are either absent or vestigial as
in submerged hydrophytes.
Notes:
@ Photoactive stomata: In most mesophytes, stomata open during day and
close during night.
@ Scotoactive stomata: In fleshy xerophytes (succulents or malacophytes),
the stomata open during
@ night hours. e.g. Agave, Aloe, Opuntia, Bryophyllum etc.
Mechanism of opening and closing of stomata

@ Shape of the guard cell is the most important factor for opening and closing
of stomata.
@ Turgidity of guard cells leads ………………………
@ Flaccidity of guard cells leads ………………………….
@ Stomata are called ‘turgor operative walls’.

Different theories of stomatal movement are given as:

1. Starch-Sugar Interconversion Theory
@ Proposed by Lloyd and elaborated by Sayre and Steward.
@ According to this hypothesis, turgor changes in guard cells are brought about
by reversible enzymic interconversion of starch into sugar under different
condition as follows.
pH high-7 , Day
Starch +iP Sugar
pH low- 5, Night
Starch
STOMATA CLOSE pH 5.0 pH 7.0 STOMATA OPEN
GLUCOSE-1-PHOSPATE
Hexokinase & ATP Glucose-6-Phospate
Phosphatase
O2 Respiration
Glucose + iP
Ojections to this theory because:

• ……………………………………………………….
• ……………………………………………………….
• ……………………………………………………….
• ……………………………………………………….
2. Potassium Malate Theory/ Active K+ exchange mechanism/

Hormonal regulation theory/ Malate switch theory:
@ Given by Fujino & later modified by Levitt.
@ A rise in pH during day time causes hydrolysis of starch to form organic acid,
i.e., phosphoenol pyruvate.
@ Phosphoenol pyruvate combines with available CO2 to produce malic acid.

Malic acid dissociates to produce H+ and malate.
@ The protons (H)+ are pumped out from the guard cells into the subsidiary
cells and in the exchange, K+ ions are pumped into them, from the adjustment
subsidiary cells.
@ The accumulation of large amount of K+ in guard cells is electrically
balanced by the uptake of negatively charged ions chloride and malate with
the help of cAMP, ATP and cytokinins.
@ K+ ions immediately combine with malate and increase the osmotic
concentration of the guard cells.
@ As a result, guard cells absorb water from the nearby epidermal
(subsidiary) cells, swell up and lead to opening of stomata.
@ Reverse situation prevails during dark when stomata are closed. There is no
accumulation of K+ ions in guard cells in dark.
• Evidences:
1. The guard cells of the opening stomata contain plenty of K+ ions.

2. During stomatal opening, starch is charged to organic acids.

Scotoactive stomata (Stomatal opening in succulents)

@ In succulents like Opuntia, stomata open during night (darkness) and are
closed during the day (light). The mechanism of their opening and closing
was explained by Nishida.
@ In night there is incomplete oxidation of carbohydrates leading to the
accumulation of malic acid without release of CO2.
Night time
2C6H12O6 + 3O2 ¾¾¾¾¾→ 3C4H6O5 + 3H2O
Malic acid
@ During daytime accumulated organic acids breakdown rapidly releasing
excess CO2 keeping stomata closed.
Night time
C4H6O5 + 3O2 ¾¾¾¾¾→ Pyruvic acid + CO2
QUICK DIGEST:
@ Active K+ exchange theory is the most accepted theory on mechanism of
opening and closing of stomata. [IOM 1997]
@ Opening of stomata is due to the influx of K+ into guard cell and closing
of stomata is due to the efflux of K+ into guard cell.
@ The extent of K+ in guard cells determines the size of the stomatal
opening.
@ At the time of opening of stomata, the turgidity of guard cells is maintained
by K+, Cl- and malate ions.
@ Primary osmolite involved in the opening of stomata is Potassium malate.
[NEET 2008]
@ Opening of stomata is active process while closing of stomata is a passive
process.
@ Maximum opening of stomata occurs in blue light, followed by red light and
no opening occurs in green light, UV-light and far red light.
@ Action spectrum for transpiration is blue red.
@ Rate of transpiration is measured by Potometer. [IOM 2006]
@ Principle of all potometers is that absorption is proportional to

transpiration.
@ Degree of opening of stomata or stomatal pore is measured with the help
of instrument called Porometer.
Factors Affecting Stomatal Movement
1. Light: action spectrum for transpiration is Blue>Red. Blue light .
2. Temperature: Usually, an increase in temperature results in increased
stomatal opening provided that water does not become a limiting factor.
3. CO2 concentration: Reduced CO2 concentration favours opening of stomata,
while increase in CO2 concentration promotes closing of stomata. [NEET
2018]
4. Water deficit: Water deficit in leaves causes stomatal closure due to rise
in DPD of epidermal cells.
5. Growth hormone: Cytokinins are essential for the opening of stomata, while
abscisic acid takes part in closing of stomata.
6. Atmospheric humidity: In humid environment, stomata remain open for longer
period while in dry environment they remain closed for longer period.
7. pH: High pH is required for the opening of stomata, while low pH induces
closure of stomata.
8. Mechanical shocks: Causes closing of stomata.
9. Leaf area: It is proportional to the rate of transpiration.
Factors Affecting the Rate of Transpiration

A. External Factors (R. H.) of atmosphere.
1. Relative Humidity: When relative humidity is high, the rate of
transpiration decreases. When relative humidity is low and the air is not
saturated with moisture, the rate of transpiration increases.
• Rate of transpiration is inversely proportional to relative
humidity.
• Most important factor for regulation of transpiration is R. H.
• Stomata remain widely open even at high R.H, but rate of
transpiration is low at high R.H.
2. Temperature: Transpiration increases with the increases in temperature.
3. Light: Maximum opening of stomata occurs in blue light, followed by red
light and no opening occurs in green light, UV-light and far red light.
4.Wind:
• Rate of transpiration is increased in gently blowing wind.
• Rate of transpiration is decreases in violently blowing wind
5. Available soil water: If available water in the soil is low is low, the
transpiration is also low.
6. Atmospheric pressure: Rate of transpiration is inversely proportional to
the atmospheric pressure.
B. Internal Factors
1. Leaf surface area: In general more is the leaf area, more will be the
transpiration.
2. Root/shoot ratio: An increase in the root/shoot ratio causes an increase
in the rate of transpiration.
3.During water stress in plants, abscisic acid (ABA) is produced that causes
closure of stomata.
Antitranspirants
@ Substances which when applied stops or reduce the rate of transpiration
without affecting normal gaseous exchange.
a. Natural anti-transpirant: ……………………………….
• ……………………………………………………….
• ……………………………………………………….
[IOM, MOE, BPKIHS]
b. Synthetic antitranspirants: ………………………………
• ……………………………………………………….
@ Transpiration from the two leaf surfaces are compared by dry paper strips
previously soaked in solution of cobalt chloride
• Dry or anhydrous cobalt chloride paper strip: Blue
• Hydrated cobalt chloride paper strip: Pink. @ Pink colour is due to Pani
@ Transpiration rate of a dorsiventral leaf is higher on the lower surface

due to higher number of stomata. [BPKIHS]
Wilting
@ Wilting occurs when transpiration > water absorption due to loss of
turgidity. It occurs when xylem is blocked.
@ Wilting is of 3 types:
a. Incipient wilting: Initial phase, not visible
b. Temporary wilting: Wilting of leaves at noon and their recovery towards
evening .
c. Permanent wilting: Plant fails to regain their original state
@ During wilting, the sequence of events shall be:
Exosmosis → Plasmolysis → Temporary wilting → Permanent wilting
Guttation
@ Guttation is loss of water from uninjured parts or margin of leaves in the
form of water droplets.
@ Guttation takes place through …………………………….
@ Hydathodes are surrounded by …………………...
@ Guttation occurs during ………………………………………..
@ Guttation is not due to activity of hydathodes but due to root pressure.

@ Guttation occurs in herbaceous mesophytes like Colocasia, Oat, Tomato,
Barley etc.
QUICK DIGEST:
@ Root pressure is the cause of guttation.
@ Hydathodes remain always open.
@ Lost water in guttation is impure as it may contain dissolved minerals.
Figure: T.S. of leaf margin showing hyadathode

@ Main differences between transpiration and guttation are:
Transpiration Guttation
Water is lost in the form of water Water is lost in the form of liquid
vapour form droplets.
It occurs during day time During night or early morning
Released water is 100% pure Released water is impure
Occurs through stomata or Occurs through hydathodes

epidermis or cuticle or lenticels
It is a controlled phenomenon It is an uncontrolled phenomenon
Exudation
@ Exudation is the loss of water sap from incision of plant body parts.
@ Exudation of water from leaf incision: Bleeding. [IE]
@ Exudation of water in the form of liquid droplets from leaf margins:

Guttation.
Advantages of Transpiration
@ It plays an important role in the upward movement of water, i.e., ascent
of sap.
@ It helps in absorption and translocation of mineral salts.
@ Rapid evaporation of water from the aerial parts of the plant through
transpiration brings down their temperature. Thus, it prevents them from
excessive heating. This is also known as cooling effect.

Plant Water Relations

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By Dr Kabir Pokhrel C/B online class 2022/23

PLANT WATER RELATIONS

@ Father of plant physiology……………………………………………

@ Father of Indian plant physiology: Sir J.C. Bose

1. Short distance (Cell to cell transport) and

@ Short-distance (Cell to cell transport):

• Diffusion is obvious in gases and liquids.

and relative density.

• In transpiration, water vapour from intercellular spaces diffuses into

@ Channel Protein : Channel protein forms a channel or tunnel in the cell

1. Uniport: molecule of a single type move across a membrane

Diffusion pressure and diffusion pressure deficit (DPD)

@ Diffusion pressure deficit (DPD) is the index of absorbing power of a

DPD µ Concentration of the solution

So the order of increasing DPD will be:

2. Hypertonic solution: ………………………………………………..

Figure: Effect of different types of solution on plant and animal cell

Figure: Types of membrane & movement of substance across different

1. Permeable membranes: Such membranes or walls allow diffusion of both,

Figure: Demonstration of Osmosis

Osmotic pressure (O.P.)

@ Highest O. P. is found in halophytes and minimum O.P. is found in

@ The osmotic pressure of a solution is measured by ……………….

@ O.P. of cell is measured by incipient plasmolysis.

Concept of Water Potential:

i.e., Water potential = -D.P.D.

Turgor pressure (T.P.) and Wall pressure (W.P.)

• Turgor pressure is a hydrostatic pressure with which cell sap presses

………………………………………….. due expanding protoplasm.

Figure: Cell showing T.P., W.P. and O.P.

DPD for fully turgid cell:

Therefore in a fully turgid cell:

@ In normal condition OP > TP

@ Bacteria cannot survive in salted pickles due to plasmolysis

@ When a cell is plasmolysed after being kept in a hypertonic solution, a

solution, it is called ……………………..

phenomenon of ………………………... [BPKIHS 2006]

@ Agar-agar is a very efficient imbibant. It imbibes 99 times its mass of water

@ Agar-agar is a phyco-colloid obtained from red algae: Gallidium,

Absorption of Water by Plants:

@ The soil transported by wind is referred to as Eolian soil and those

• Sandy soil (85% sand + 15% clay or silt or both)

@ Loamy soil is considered best for cultivation of crop as it maintains an

Pathway of Water Movement in Plants

Metaxylem Protoxylem Pericycle cells Endodermis

@ Pathway followed by water is of two types, i.e., symplast pathway and

without crossing any membranes. The apoplast is the continuous system

the plasmodesmata. The symplast consists of an entire network of cell

Epiblema Cortex Endodermis Xylem

Figure: Pathway of water movement

Mechanism of water absorption:

a. Active absorption through osmotic mechanism:

b. Active absorption through non-osmotic mechanism:

Differences between Active and Passive absorption

Factors affecting water absorption by plants:

concentration of salts. Such plants cells have high osmotic concentration

@ Best experiment to show path of ascent of sap is Ringing or Girdling

the tissues outer to vascular cambium).

@ In this experiment; all tissue

@ According to this theories, the forces required for ascent of sap is