Lecture 4

Translocation of water - Xylem Transport

 Translocation of water - Xylem Transport

 Otherwise called as “Ascent of Sap”

 Upward movement of water in plant - “Ascent of Sap”

 Sap moves through xylem – Xylem Sap

 Direction of translocation of Sap – Normally upward

 Unidirectional Movement
 Xylem Transport
Xylem

1. Water conducting Tissue – Complex in nature

2. Kinds of cells – Tracheids & Vessels

3. Dead Cells

4. Angiosperms - Vessels & Tracheids

5. Gymnosperms – Tracheids

V T
The tracheids and vessel elements are dead, specialized to
conduct water from root

Vessels
•Thickened walls

•Thickenings are due to lignin

•Vessels are arranged end to end

•End wall is either partially or fully dissolved

•Perforated side wall

Tracheids

•Tracheids are elongated dead cells with sloping end walls Pits

with pits

•The walls are thickened with a material called lignin

 Mechanism of Ascent of Sap

A. Vital theory

B. Root pressure theory

C. Physical force theory

D. Transpiration Pull Theory

A. Vital theories
• According to vital theories, the ascent of sap is under the
control of vital activities in the stem.

• According to Godlewski (1884) – Ascent of sap takes

place due to the pumping activity xylem tissues

• According to Bose (1923) – Upward translocation of

water takes place due to pulsatory activity of the living
cells of the inner most cortical layer just outside the
endodermis.
 B. Root Pressure Theory

It was first observed by Stephen Hales

A pressure is developed in the tracheids of xylem of root

in fully turgid condition which can raise the water to a

Certain height in the xylem

This pressure is called as root pressure

 Root Pressure

Exudation of water from the cut stem when the shoot is

removed near the soil level

Root pressure is responsible for Ascent of sap in plants

 Objections of RP Theory

1. RP can raise the water up to 21 meters; Tall tree?

2. RP lowest during summer
3. High during winter
4. RP is very little in Red Wood
C. Physical force theories
Various physical forces may be involved in Ascent of sap.

1. Atmospheric pressure
• This does not seem to be convincing because it cannot act on water
present in xylem in roots. In case it is working and then also it will not
be able to raise water beyond 34 feet .
• 2. Imbibition
• Sachs (1878) supported the view that ascent of sap could take place
by imbibition through the walls of xylem. But imbibitional force is
insignificant
• 3. Capillary force
• In plants the xylem vessels are placed one above the other forming a
sort of continuous channel which can be compared with long capillary
tubes and it was thought that as water rises in capillary tube due to
capillary force in the same manner ascent of sap takes place in the
xylem.
 Transpiration Pull Theory

• Cohesion & Adhesion Theory

• Hydrostatic Tension Theory

• Proposed by Dixon & Jolly

• Based on cohesive & adhesive properties of water

ohesion: Attraction between like molecules

dhesion: Attraction between different molecules

• Water is lost from the surface of the mesophyll cells of the leaves. The
water escapes as water vapour into the intercellular spaces and finally
into the atmosphere through stomata. The sequence of events are

   Water vapours diffuse out from the intercellular space to the

atmosphere.

   Evaporation of water vapours from thin walled mesophyll cells (leaf)

into the intercellular space.

   Reduced water content in the cell

   Increased osmotic potential  in cell sap

   Movement of water from adjacent cell by osmosis

   Movement of water from xylem to adjacent cell

• A kind of water potential gradient exists from the leaf to the root and
creates a transpiration pull.

• The water molecules have a strong force of cohesion (a tendency to

stick together). The force of cohesion is even more in narrow tubes
such as xylem vessels.

• In addition, there is also a force of adhesion between the walls of

xylem vessels and water. The ligno-cellulose walls not only provide
strength but have a strong affinity for water molecules.

• Both the forces, adhesion and cohesion together, help to form thin,
unbroken columns of water in the capillaries of xylem vessels. The
transpiration pull lifts the water column up by creating a tension.

• It is almost similar to the liquid being taken up from a straw.

 Transpiration
Adhesion

Transport

Absorption Cohesion
 Lecture 5
Transpiration
Loss of water in the form of water vapour from
aerial parts of plant

Through the plant

About 97% of absorbed water lost by transpiration

 Types of Transpiration

1. Stomatal Transpiration – 90 to 95%

2. Cuticular Transpiration – 5 to 10%

3. Lenticular Transpiration – 0.1%

Cuticular transpiration

The loss of water through the

impervious cuticle – 5 to 10%

Lenticular transpiration

The loss of water through the

lenticels of woody stems. 0.1%
of the total transpiration loss
of water
 Stomatal Transpiration
 Transpiration through Stomata
 Most of the transpiration takes place through
stomata - 90-95%
 Structure of Stomata
a.Dumbbell shape b. Kidney shape

• Two distinct types of guard cells

exist, kidney shaped and dumb-
bell shaped.
• Kidney-shaped guard cells (a) are
found in dicotyledons whereas
dumb-bell-shaped guard cells are
found in grasses ( b).
• Dumbbell shaped guard cells are
more advanced in evolutionary
terms and more efficient
physiologically because guard
cells of grasses require fewer
solutes and less water to achieve
a given unit increase in aperture
 Stomata are minute pores of
Structure of stomata elliptical shape surrounded by two
specialized epidermal cells known
as guard cells.

The guard cells are kidney shaped

in dicots and dumbbell shaped in
monocots.

The portion of guard cell wall lying

close to stomatal aperture is thick
and inelastic.

The outer cell wall is thin, elastic

and permeable.
Each guard cell has a cytoplasm lining
and a central vacuole containing cell
sap. Its cytoplasm contains a nucleus
and a number of chloroplasts.
Thin &
elastic wall The epidermal cells surrounding guard
cells are specialized and are known as
subsidiary cells. They support the
movement of guard cells. The size,
shape and number of stomata of
guard cells vary from plant to plant.
 Types of Stomata
1. Hypostomatus – Only on lower surface

2. Epistomatus – Only on upper surface

3. Amphistomatus – Present both the surfaces

4. Anisostomatous – More on lower surfaces

5. Isostomatus – Equal on both surfaces

6. Potamogeton – Absent or functionless

Stomata are usually confined in more
numbers on the lower sides of the
Leaves

In most of the monocots, they are

equally distributed on both sides of
leaves

In aquatic plants, stomata are present

only on the upper surface of the
floating leaves
 MECHANISM OF STOMATAL TRANSPIRATION

3 steps in the Mechanism

1. Osmosis of water 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 vapour from intercellular

spaces to

outer atmosphere
3 steps in the Mechanism of stomatal Transpiration
 OSMOTIC DIFFUSION IN TRANSPIRATION

 Inside the leaf, the mesophyll cells are in contact with xylem
and intercellular spaces above the stomata

 When mesophyll cells draw water from the xylem they

become turgid and high water potential by osmosis

 Water is released in the form of vapour into intercellular

Spaces close to stomata by osmosis

 Stomatal movement : Water pass to the GC in the form of

water vapour and then to atmosphere through stoma
( Pore) by simple diffusion.

 Now, the water potential of mesophyll cells become low and

hence, they draw water form xylem
 GC Turgid GC Flaccid
Stomata open Stomata closed

Day time – Stomata Open

Night time – Stomata Closed
 Theories for mechanism of
opening and closing of stomata

1. Theory of Photosynthesis

2. Starch – Sugar Inter conversion Theory – Steward Theory

3. Theory of Proton Transport and Hormonal Regulation

 Theory of Photosynthesis
GC contains chloroplast not other epidermal cell

resence of light photosynthesis takes place in GC

Production of sugars in GC

Water potential decreases in GC

Water enters in to GC

GC become Turgid Stomata Open

 Starch – Sugar Interconversion Theory - Classical Theory

Based on effect of pH on Starch Phosphorylase enzyme

Light & High pH

Glucose -1-PO4
Starch + Pi
(Soluble)
(Insoluble)
Dark & Low pH

Water potential ↑ in GC Water potential ↓ in GC

Water comes out from GC Water enter into GC

GC become Flaccid GC become Turgid

Stomata Closed Stomata Open

 Steward Theory of Stomatal opening & closing

Stomata
closed Starch Pi

Dark & pH 5 Starch Phosphorylase

Light & pH 7
Glucose -1-PO4

Phosphoglucomutase
ATP
Glucose -6-PO4
Hexo Phosphatase
kinase
Glucose + Pi

Stomata
 Proton (H+) – K+ Exchange Pump

Reaction in GC Reaction in GC
during Day during Night
H+
No
Photosynthesis
Photosynthesis
K+ No H+
K+
Malic Acid Cl-
C3H5O3COOH Cl-
Water potential ↑
C3H5O3COO- + H+
K+ & Cl- Act as Osmolytes
Water
comes out
Stomata Open
Water potential ↓ in GC
Flaccid

GC become Turgid Water enter into GC Stomata

Closed
 Significance of Transpiration
Advantages - Necessary

1. Role in movement of water

2. Absorption and Translocation of Nutrients

3. Regulation of plant Temperature

Disadvantages - Evil

 95 – 97 % applied water lost

 Only 1% utilized for metabolic activities

 Unnecessary wastage of water & Energy

 Cannot avoid Transpiration

Because
Stomata should be open for Gaseous exchange
for Photosynthesis & Respiration

1% Water is required for metabolic activity

Transpiration – Necessary Evil

 Guttation
Exudation of liquid water droplets from the
uninjured margins and tips of the leaves where
vein ends through special type of cell –
Hydathode or water stomata

Takes place usually early in the morning when

the rate of absorption and root pressure are
high while the transpiration is very low

Example plants

Tomato, Strawberry,
Colocasia, Nasturtium
and grasses
Guttation in Grasses
Guttation in Strawberry leaves
 Differences between Transpiration and Guttation
Transpiration
It takes place throughout the
Day
It does not depend on root
pressure
Water is lost from aerial parts
of plants in the form of water
vapour
It occurs mostly through Stomata
and also take place through cuticle
and lenticels
Transpired water is pure
The leaf temperature is reduced
Guttation
It takes place at night and in
the early morning
Depends on root pressure
Water lost as liquid oozes out from
uninjured margins of leaves
It occurs only through Hydathodes
(water stomata)
Guttation fluid contains salts and
Sugars as aqueous solution
There is no such effect
 Antitranspirants
Chemicals or substances that are used to
reduce transpiration in plants

Three Groups of Antitranspirants

1. Stomatal Closing Type

2. Film Forming Type

3. Reflective Type
 Stomatal closing type
 These antitranspirants induce the closure of
stomata and reduce the rate of transpiration
 The rate of CO2 diffusion into the leaf is also
reduced leading to low photosynthetic rate
1. Phenyl Mercuric Acetate
(PMA)
2. Abscisic Acid (ABA)
3. Carbon-di-Oxide (CO2)

Low concentration of PMA & CO2 promising

results by partial closure of stomatal pore
 Film forming type

This type forms a thin film coating on the

surface of leaf and inhibits the loss of water
vapour from the leaf

They also inhibit (minimum) CO2 entry

E.g. Waxes, Plastic films, Silicone oils

 Reflective type

This type of chemicals increases the light

reflection by the leaves, thus decreasing the
leaf temperature

The water loss is reduced without affecting the

CO2 assimilation

E.g. Kaolinite (Kaolin), Lime water (Lime wash)

 Factors affecting transpiration
1. Light
• Plants transpire more rapidly in the light than in the
dark.
• Light stimulates the opening of the stomata and speeds
up transpiration by warming the leaf.
2. Temperature
Temp increases Transpiration increases
(30°C three times faster than 20°C )
3. Humidity : Less and leads to decrease transpiration
4. Wind : no wind causes reduced transpiration
5. Soil water : Transpiration is more with water availability

one acre of corn plants may transpire 400,000 gallons of

water in a growing season
 Review
1.Mention the different type of transpiration?
2.Write the significance of transpiration?
2.Write the three type of anti transpirants with example
3.Match the following

Guttation ATP requirement &

Glucose

Starch-sugar Stomata
interconversion theory
Steward theory K&H
Proton transport theory pH based & G-1-P

Transpiration Hydathode
 Review
1.Mention the different type of transpiration?
a.Stomatal Transpiration – 90 to 95%
b. Cuticular Transpiration – 5 to 10%
c.Lenticular Transpiration – 0.1%

2.Write the significance of transpiration?

1. Role in movement of water
2. Absorption and Translocation of Nutrients
3. Regulation of plant Temperature

3.Write the three type of anti transpirants with example

• Stomatal Closing Type : PMA, CO2 and ABA
• Film Forming Type : Waxes, Plastic films, Silicone oils
• Reflective Type : Kaolinite (Kaolin), Lime water (Lime wash)
 3.Match the following

Guttation Hydathode

Starch-sugar pH based & G-1-P

interconversion
theory

Steward theory ATP requirement

& Glucose

Proton transport K&H

theory

Transpiration Stomata