Stomatal Opening and Closing
Stomata are functional unit of the epidermis serving the exchange of gases between
the intercellular spaces of the plant and its surrounding. They are especially common
and of characteristic shape at the epidermis of the leaf's underside of most species.
Their development differs from plant group to plant group, but unequal cell divisions
are always involved. A functional unit consists of the guard cells themselves that
contain nearly always chloroplasts and of their neighbouring subsidiary cells that are
usually devoid of chloroplasts.
Comment
Mechanism and Regulation of Stomata Movements
The exchange of oxygen and carbon dioxide in the leaf (as well as the loss of water
vapour in transpiration) occurs through pores or stomata (singular = stoma).
Normally stomata open when the light strikes the leaf in the morning and close during
the night.
The immediate cause is a change in the turgor of the guard cells. The inner wall of
each guard cell is thick and elastic. When turgor develops within the two guard cells
flanking each stoma, the thin outer walls bulge out and force the inner walls into a
crescent shape. This opens the stoma. When the guard cells lose turgor, the elastic
inner walls regain their original shape and the stoma closes.
Comment
Factors affecting Stomatal Movement
Water
Guard cells can emit water in three different directions:
1. Outwards
2. Into the neighbouring subsidiary cells
3. Into the respiratory cavity that is a part of the intercellular system lying
beneath the guard cells.
An equilibrium between the water vapour in the atmosphere and the respiratory cavity,
results when the stomata are opened. Plants form an intermediate distributor, since a
large difference in water potential between the moist soil and the normally dry
atmosphere is very common. Plants profit from the concentration gradient (that is a
gain of energy for them), while the closing movements of the stomata exert a decisive,
regulating influence. They close when too much water is lost or when not enough
supply exists. The osmotic pressure of the stomata is far larger in the guard cells than
in the subsidiary cells. This ratio shifts in favour of the subsidiary cells when the
stomata are closed.
Light and Carbon Dioxide
The stomata of most plant species are closed in darkness. Light stimulates opening.
The action spectrum is similar to that of photosynthesis. Blue light is especially
effective. The stomata of CAM-plants, like Crassulaceans, are open during the night.
They depend on the accumulation of carbon dioxide during the night. These plants
store the carbon dioxide as malate or aspartate and feed it into the Calvin cycle during
daytime. Open stomata would cause intolerable transpiration losses in the areas that
CAM-plants live.
A low concentration of carbon dioxide (in the respiratory cavity) causes the stomata to
open. A high concentration leads to their closing. Photosynthesis starts with the first
light of the day, because enough carbon dioxide has been accumulated.
Photosynthesis takes place in guard cells too, since they contain chloroplasts in
contrast to the subsidiary cells. This activity again is related to the rise of the osmotic
value and thus also to the opening of the stomata.
StomatalOpeningandClosing
Controlling cycles regulating the movements of stomata. A guard cell is depicted
schematically.
On what is the rise or lowering of the osmotic value based? Good evidence exists that
light exerts its effect mainly by decreasing the intercellular and intracellular
concentration of carbon dioxide. Carbon dioxide is faster consumed than supplied due
to the process of photosynthesis. This causes an intercellular deficit close to the
stomata's opening. The photosynthetic activity within the guard cells leads to a
decrease of the intracellular level of carbon dioxide and this causes water to be drawn
from the subsidiary cells.
It turned out that the water-uptake is preceded by an uptake of potassium ions.
Potassium ions are actively pumped (by a potassium pump) from the subsidiary cells
into the vacuoles of the guard cells. At the same time, anions (chloride, malate)
accumulate within the vacuoles. Protons are given off to the subsidiary cells.
The actual importance of the potassium pump for the guard cell movement is best
demonstrated with a fungal toxin called Fusicoccin (from the fungus Fusicoccum
amygdali). This toxin activates the potassium pump. If consequently, the toxin is
applied to the stomata, then the loss of water becomes higher than its supply,
resulting in withering. The biological advantage for the fungus lies in the open stomata
since they are, beside wounds, the only places where its hyphes can penetrate the leaf
tissue. The total water balance is hardly or not at all influenced, as long as the effect
of Fusicoccin remains a local one. In principle, one open stomata is enough for a hype.
Temperature
Stomata tend to open more with an increase in temperature and close with a decrease
in temperature. In some plant species, stomata remain closed even under continuous
light at OoC. However, if the temperature is increased, stomatal opening in these
StomatalOpeningandClosing
species increases. At temperatures higher than 30oC there is a decline in stomatal
opening in some species.
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