Stomatal Regulation

 Stomata are the tiny openings present on the epidermis of leaves which aid in gaseous
exchange between plants and surrounding. They are found only in the aerial parts of the
plants.
 The stomatal opening and the guard cells are together referred as stoma.
 The number of stomata present on the surface of leaves varies from species to species.
 The number of stomata present in the lower surface is always greater than that in upper
surface. However, in many cases stomata are absent on the upper surface, for example in
xerophytes.
 Stomata are scattered in case of dicot leaves while they are arranged in parallel rows in
case of monocot leaves.
 The kidney shaped cells present around the stomata are called guard cells. They are living
and consist of chloroplasts and a large amount of cytoplasm.
 The accessory or subsidiary cells refers to the epidermal cells present around the guard
cells.
Opening and Closing of Stomata
 The opening and closing of stomata depends on the presence of sugar and starch in the
guard cells.
 During the day time, photosynthesis takes place in the chloroplasts and so guard cells
contain soluble sugar which increases the sap concentration of guard cells. So, water
from surrounding cells move towards guard cells through osmosis and they become
turgid. Thus, stomata are opened.
 During the night time, there is no sunlight and so, no photosynthesis takes place and the
sugar present in the cells is converted into insoluble starch. The sap concentration of the
guard cells is low and so water moves from guard cells to the surrounding cells making
the cell flaccid and thus, stomata are closed.
 The starch-sugar conversion depends on the pH of the cell. At night, no photosynthesis
takes place so, carbon dioxide accumulates in the cell, makes acidic conditions and sugar
is converted into starch.
 During day time, carbon dioxide is consumed in the photosynthesis so basic condition is
created and starch is converted into sugar.
 Bonner and Galston Hypothesis
 Bonner and Galston proposed that the mechanism of opening and closing of stomata is
independent of the presence or absence of sunlight and depends on the carbon dioxide
concentration of stomatal cell.
 When concentration of carbon dioxide equals the concentration of carbon dioxide i.e.
0.03% then guard cells are flaccid and stomata are closed.
 As the density of carbon dioxide decreases the stomata starts to open when concentration
of carbon dioxide becomes 0.01% stomata are fully opened.
 During day time, concentration of carbon dioxide decreases below 0.03% because of its
consumption in photosynthesis so the stomata are opened.
 During night time, concentration of carbon dioxide is 0.03% because of it is not
consumed in photosynthesis so guard cells are flaccid and the stomata are closed.
Active Potassium (K+) Theory
 This theory is the universally accepted theory for the mechanism of opening and closing
of stomata.
 According to this theory potassium ions are responsible for opening and closing of
stomata. This was first observed by Fujino in 1967 that the stomata open due to transfer
of potassium ions from the surrounding cells into the guard cells.
 Uptake of potassium ions increases the osmotic concentration of guard cells and so, water
from surrounding cells enters into the guard cell which leads to opening of stomata. So,
uptake of potassium ions controls the water potential gradient.
 In 1974, Levitt found that uptake of protons by the guard cells with the help of ATP
increases the pH of guard cells which results into conversion of starch into organic acid
like malic acid.
 During the day time, in presence of light, starch is converted into malic acid which
dissociates into malate ions and protons(H+).
 Protons are exchanged with the potassium ions by the guard cells. This exchange requires
ATP. So, the concentration of potassium ions increases and that of protons decreases
thus, pH increases.
 The potassiums ions taken by the cell is balanced by chloride ions and malate ions,
produced after hydrolysis of starch into malic acid.
 The potassium ions form potassium malate which enters in the cell sap. Water potential
decreases and the osmotic concentration of the cell decreases, endoosmosis takes place,
guard cells become turgid and the stomata opens.
 During night, there is no light so concentration of malic acid is low. Potassium ions move
out of cell and protons enters the cell and pH of the cell decreases. Under these
conditions, exosmosis takes place, cells become flaccid and stomata closes.
 Factors Affecting Stomatal Movement
1. Light- Stomata open in light and close in absence of light. Optimum amount of light
required is different for different species.
2. Temperature- Generally, stomatal opening increases with increase in temperature if
water is not limiting factor.
3. When availability of water is high stomata are opened as the guard cells are turgid and
when availability of water is low stomata are closed as the guard cells are flaccid.
4. Carbon dioxide concentration- Low CO2 concentration causes opening of stomata
while an increase in CO2 concentration causes closing of stomata.