Summary
Regulation of gas exchange by stomata of adjustable aperture size in the leaf epidermis is crucial to a plant’s physiology: sufficient CO2 must be admitted into the leaves for growth, but loss of water, usually the limiting resource for terrestrial plants, must be minimized. In essence, the study of stomatal movements is an inquiry into accumulation and dissipation of K + salts, which account for the bulk of the osmotic change associated with stomatal movements.
The cardinal event in stomatal opening is the activation of the H +-extruding ATPase of the guard cell plasmalemma, which hyperpolarizes the membrane, strengthening the driving force for passive K> + inward permeation and opening the K +-specific inward-rectifying channel. Ultimately, K + is transported into the vacuole, implying the importance of less well studied ion traffic across the tonoplast. Depolarization, in order to develop an outward driving force for K +, is required for stomatal closure and results from anion efflux, which is stimulated by Ca 2+, the role of which is not fully understood.
Abscisic acid diminishes gas exchange by activating the K +-out channel in a Ca 2+-independent fashion. In addition, by stimulating a plasmalemma Ca 2+-in channel or by activating aphosphoinositide signal transduction pathway, it elevates cytosolic Ca 2+ concentration. In turn, elevated Ca 2+ concentration shifts more negative the gating potential of the K +-in channel and activates the anion efflux channels. Long term effects of abscisic acid on guard cells have not been investigated, but its integrating function as a long distance messenger is currently under intense investigation.
Potassium concentration fluctuations do not occur in isolation. If stoichiometric, accumulation of Cl − by an unknown active transport process would balance K +, and the pH within the cell would be stabilized even as protons were pumped out, however, Cl − accumulation is insufficient. Synthesis of malate, during which two protons are released to solution, ‘replenishes’ extruded protons not accounted for by Cl − uptake. These and other processes ancillary to K + fluxes are addressed here.
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Outlaw, W.H., Zhang, S., Hite, D.R.C., Thistle, A.B. (1996). Stomata: Biophysical and Biochemical Aspects. In: Baker, N.R. (eds) Photosynthesis and the Environment. Advances in Photosynthesis and Respiration, vol 5. Springer, Dordrecht. https://doi.org/10.1007/0-306-48135-9_9
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