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Explain the mechanism of opening and closing of stomata.

Solution
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When the guard cells surrounding the stomatal pore are in a turgid state, stomata open and when the guard cells become flaccid stomatal opening gets closed.
Mechanism of Opening and Closing of Stomata:
  • Opening and closing of stomata may be called stomatal movement.
  • Several hypotheses have been proposed to explain the stomatal movement.
1. Starch:Sugar hypothesis:
This hypothesis was proposed by J.D. Sayre (1923).
  • Prior to Sayre, Lloyd (1908) showed the presence of starch in the guard cells.
  • According to starch-sugar hypothesis during day time, $$CO_2$$ is used in guards cells in photosynthesis and pH of guard cells increases (7.0).
  • At this pH, the phosphorylase enzyme becomes active in guard cells and starch is converted into glucose-1-phosphate.
  • On account of this, DPD of guard cells increases and they become turgid due to endosmosis and stomata open.
  • During the night, photosynthesis stops and $$CO_2$$ accumulates in the guard cells, due to which the pH of guard cells decreases (5.0).
  • At this pH, glucose-1-phosphate converts into starch.
  • DPD of guard cells decreases and they become flaccid due to exosmosis, causing the closing of stomata.
Starch + iP → Glucose - 1 - phosphate
2. Steward's hypothesis:
  • According to this hypothesis, an increase in pH of guard cells results in the conversion of starch in glucose-1-phosphate, and then into glucose-6-phosphate which finally converts into glucose.
  • Glucose and glucose-6-phosphate are more soluble in water as compared to glucose-1-phosphate.
  • This results in an increase, in the osmotic concentration of guard cells leading to the opening of stomata.
  • Biochemical reaction during day time:
Starch + iP → Glucose-1-phosphate
Glucose-1-phosphate → Glucose-6-phosphate
Glucose-6-phosphate → Glucose + iP
  • Biochemical reactions during night time:
Glucose + ATP → Glucose-1-Phosphate
Glucose-1-Phosphate → Starch + iP
3. Active Potassium Ion Transport theory:
  • This theory was proposed by Japanese scientists. Imamura and Fujino (1959) and was modified by Levitt (1974).
  • Presently this is the most accepted theory for explaining the stomatal movement.
  • According to this theory, during the presence of light malic acid is formed in guard cells which breaks into malate and in hydrogen ion ($$H^+$$).
  • These $$H^+$$ move out of guard cells and pass to subsidiary cells (influx) or epidermal cells.
  • $$K^+$$ from subsidiary cells or epidermal cells move into guard cells where they react with the malate and form potassium malate.
  • This increases the osmotic concentration of the guard cells.
  • Increase in osmotic concentration of guard cells results into endosmosis in these cells and they become turgid causing opening of stomata.
  • During night efflux of $$K^+$$ from guard cells to adjacent epidermal cells takes place.
  • $$H^+$$ flow from epidermal cells into guard cells where they react with malate to form malic acid.
  • This principle is also called as $$K^+$$ ion exchange or anion exchange or photoactive proton transport theory.
  • At present this theory is the most accepted theory for explaining the opening and closing of stomata.

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