Calcium-based signalling systems in guard cells

Calcium is a ubiquitous intracellular signal responsible for controlling numerous cellular processes in both plants and animals. As an example, Ca²⁺ has been shown to be a second messenger in the signal transduction pathways by which stomatal guard cells respond to external stimuli. Regulated increases in the cytosolic concentration of free calcium ions ([Ca²⁺ ]_cyt ) in guard cells have been observed to be a common intermediate in many of the pathways leading to either opening or closing of the stomatal pore. This observation has prompted investigations into how specificity is encoded in the Ca²⁺ signal. It has been suggested that the key to generating stimulus-specific calcium signatures lies in the ability to access differentially the cellular machinery controlling calcium influx and release from intracellular stores. Several important components of the calcium-based signalling pathways have been identified in guard cells including cADPR, phospholipase C-InsP₃ , InsP₆ and H₂ O₂ . These data suggest that the pathways for intracellular mobilization of Ca²⁺ are evolutionarily conserved between plants and animals. ABBREVIATIONS: ABA, abscisic acid; [Ca²⁺ ]_cyt , cytosolic free calcium concentration; [Ca²⁺ ]_ext , external calcium concentration; I_K,in ; inward-rectifying K⁺ currents; InsP₃ , inositol-1,4,5-trisphosphate; InsP₆ , inositol hexakisphosphate; PLC, phospholipase C; PLD, phospholipase D; PA, phosphatidic acid; H₂ O₂ , hydrogen peroxide; AAPK, ABA-activated serine-threonine protein kinase; cADPR, cyclic adenosine 5'-diphosphoribose; U73122, 1-(6-{[17â-3-methoxyestra-1,3,5(10)-trien-17-yl]amino}hexyl)-1H-pyrrole-2, 5-dione; RyR; ryanodine receptor; CICR; calcium-induced calcium-release; I_Ca , inward calcium current.