Theories of Stomatal

Movement: 4 Theories (With

Diagram)
The following four points will highlight the four important
theories of stomatal movement.
The four important theories of stomatal movement are:
(1) Theory of Photosynthesis in Guard Cells (2) Starch
Sugar Inter-conversion Theory (3) Theory of
Glycolate Metabolism and (4) Active K+ Transport or
Potassium Pump Theory and Role of Abscisic Acid
or Active Potassium Pump Theory.
Many theories have been proposed regarding opening and closing of
stomata.

The important theories of stomatal movement are as

follows:
1. Theory of photosynthesis in guard cell

2. Starch Sugar inter-conversion theory

3. Theory of glycolate metabolism

4. Active potassium transport ion concept

Theory # 1. Theory of Photosynthesis in Guard Cells:

Von Mohl (1856) observe that stomata open in light and close in the
night. He then proposed that chloroplasts present in the guard cells
photosynthesize in the presence of light resulting in the production
of carbohydrate due to which osmotic pressure of guard cells
increases.

Its explanation is based on following sequence:

Light → Photosynthesis in guard cells → Formation of sugar
Increase of osmotic pressure of cell sap → Endosmosis takes place
from subsidiary cell to guard cell → Increase of TP in guard cells →
Stomata open.
Demerits:
1. Increasing the CO2 concentration around the leaves should lead to
wide opening of stomata but here occurs their partial closure.
2. Chloroplast of guard cells are poorly developed and incapable of
performing active photosynthesis.

Theory # 2. Starch Sugar Inter-conversion Theory:

(i) According to Lloyd (1908) turgidity of guard cell depends on
inter-conversion of starch and sugar. It was supported by Loft-field
(1921). He found out that guard cells contain sugar during day time
when they are open and starch during night when they are closed.

(ii) Sayre (1926) observed that stomata open in neutral or alkaline

pH, which prevails during day time due to constant removal of
carbon-dioxide by photosynthesis. Stomata remain closed during
night when there is no photosynthesis and due to accumulation of
carbon-dioxide, carbonic acid is formed that causes the pH to be
acidic. Thus, stomatal movement is regulated by pH due to inter-
conversion of starch and sugar. Sayre concept was supported by
Scarth (1932) and Small et. al. (1942).

(iii) Yin and Tung (1948) isolated for the first time phosphorylase
enzyme from the guard cells. According to them starch is converted
into glucose-1, phosphate in the presence of this enzyme. During the
process, inorganic phosphate is also used and light and dark phases
(changing CO2 concentration) control the changes in pH. The
reaction maybe represented as follows:

(iv) Steward’s scheme:

Steward (1964) proposed another modified scheme of inter-
conversion of starch and sugar for stomatal movement. He believes
that conversion of starch to Glucose -1 phosphate is not sufficient. It
should be converted to glucose in order to increase sufficient
osmotic pressure. For this, ATP is also required which means that
the process should be through respiration in presence of oxygen.
Guard cell carries enzymes like Phosphorylase,
Phosphoglucomutase, Phosphatase and Phosphorylase. These
enzymes help in opening and closing of the stomata.

Based on the above mentioned theory, process of opening and

closing of stomata may be summarized as given below.

In Light:
Photosynthesis (1) →Decreased CO2Concentration in leaf cells (2)
→Increase in pH of guard cells(3) → Hydrolysis of starch to sugar
by enzymes (4) → Increase of O.P. of guard cells(5) → Endosmosis
of water in guard cells (6) →Increase in T.R of guard cells (7)
→Aperture opens (Fig. 4.6)

Demerits of the starch-sugar inter-conversion theory:

Many scientists do not agree with the theory of starch-sugar inter-
conversion due to the following reasons.

1. In the presence of light when starch disappears from guard cells,

malic acid appears and not the sugars.

2. Starch has not been reported in the guard cells of many monocots
such as Iris, Amatyllis, Allium.

3. According to this theory O.P. of guard cells increases due to the

formation of glucose-1- phosphate in guard cells but it is found that
the presence of phosphate ions causes the development of same O.P
as does the presence of glucose-phosphate.

4. Enzyme phosphorylase helps in conversion of starch to glucose-1-

phosphate but not in the formation of starch from glucose-1-
phosphate. This reaction is controlled by some other enzyme about
which we do not know as yet.

5. The theory could not explain the extra effectiveness of blue light
at the time of stomatal opening.

Theory # 3. Theory of Glycolate Metabolism:

Zelitch (1963) proposed that production of glycolic acid in the guard
cells is an important factor in stomatal opening. Glycolate is
produced under low concentration of CO2. He suggested that
glycolate gives rise to carbohydrate, thus raising the osmotic
pressure and also that it could participate in the production of ATP.
Which might provide energy required for the opening of stomata.
Demerits:
1. It fails to explain the opening of slomata in dark (e.g., – in
succulent plants).

2. In some plants slomata have been found to remain closed even

during daytime.

3. It fails to explain the effect of blue light on stomatal opening.

Theory # 4. Active K+ Transport or Potassium Pump Theory and

Role of Abscisic Acid:
Or
Active Potassium Pump Theory:
(Active K+ion Transport (K+ ion Pump) Mechanism)
The concept of K+ ion transport was given by Fujino. It was
supported and elaborated by Levitt & Rashke in 1975 It appears to
be an active mechanism which needs ATP. It is based on recent
observations and (explains the mechanism as follows.
A. Opening of Stomata during Daytime (in presence of
light):
Opening of stomata depends upon following conditions:
(a) Presence of light.

(b) Decrease in starch contents of guard cells.

(c) Increased concentration of malic acid in guard cells.

(d) Influx of K+ ions in guard cells.

(e) Efflux of H+ ions from guard cells.
(f) Intake of CI ions by guard cells.

(g) Low CO2 concentration in an around guard cells.

(h) High pH (more than 7) in guard cells (hence, alkaline medium of
the cell sap in guard cells).

(i) High T.P. in guard cells due to endosmosis, (turgidity of cells).

(j) TP more towards thin wall of guard cell & stomata open.

Explanation of Levitt Concept:

This is explained as follows:
In the guard cells, starch is converted into malic acid in presence of
light (during day time).

Protons (H+) thus formed are used by the guard cells for the uptake
of K+ ions (in exchange for the protons H+). This is an active ionic
exchange and requires ATP energy and cytokinin (a plant
hormone). In this way, the concentration of K+ ions increases in
guard cells. At the same time, the concentration of H+ ions
decreases in guard cells. The pH of the cell sap in guard cells also
increases simultaneously (pH becomes more than 7 and the
medium becomes alkaline).
There is also an increased uptake of CI” (anions) by the guard cells
to maintain the electrical and ionic balance inside and outside the
guard cells. The malate anions formed in the guard cells are
neutralized by the K+ ions. This results in the formation of
potassium malate.
Malate anions + K+ →Potassium malate:
Potassium malate enters the cell sap of the guard cells thereby
reducing the water potential while increasing the osmotic
concentration (and the O.P.) of the cell sap. Hence, endosmosis
occurs, guard cells become turgid and kidney-shaped and the
stomata opens.

It is also observed that the CO2 concentration is low in and around

guard ceils during day time. This is due to high photosynthetic
utilization of CO2. It helps in opening of stomata.
B. Closing of Stomata in Absence of Light (Darkness/Night
Time):
Closing of stomata depends on following conditions:
(a) Absence of light.

(b) Decreased concentration of malic acid in guard cells.

(c) Efflux of K+ ions from guard cells.

(d) Influx of H+ ions in guard cells.
(e) Acidic medium of the cell sap in guard cells.

(f) Loss of Cl– ions from guard cells.

(g) Increases CO2 concentration in and around guard cell due to
release of CO2 in respiration combined with the absence of
photosynthetic activity in dark.
(h) Presence of plant growth inhibiting hormone abscissic acid
(ABA),
(i) Loss of turgidity and loss of kidney-shape by guard cells.

All these conditions represent the reversal of the daytime events.

Under these conditions, the guard cells lose water by exosmosis and
become flaccid. This causes closing of the stomata.

Role of Plant Hormones in Stomatal Movements:

(i) Presence of Cytokinin (Plant growth regulator) is needed for the
active uptake of K+ ions

(ii) Presence of ABA (abscissic acid, a plant growth inhibiting

hormone) favours closing of stomata by blocking uptake of K+ by
guard cells in the dark. It also prevents efflux of H+ ions from guard
cells. ABA and CO2 cone, together help in lowering the pH in guard
cells and making the medium acidic. This helps in closing of
stomata. ABA act as stress hormone during drought condition.