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A central question is how specificity in cellular responses to the eukaryotic second messenger Ca 2+  is achieved. Plant guard cells, that form stomatal pores for gas exchange, provide a powerful system for in depth investigation of Ca 2+ -signaling specificity in plants. In intact guard cells, abscisic acid (ABA) enhances (primes) the Ca 2+ -sensitivity of downstream signaling events that result in activation of S-type anion channels during stomatal closure, providing a specificity mechanism in Ca 2+ -signaling. However, the underlying genetic and biochemical mechanisms remain unknown. Here we show impairment of ABA signal transduction in stomata of calcium-dependent protein kinase quadruple mutant plants. Interestingly, protein phosphatase 2Cs prevent non-specific Ca 2+ -signaling. Moreover, we demonstrate an unexpected interdependence of the Ca 2+ -dependent and Ca 2+ -independent ABA-signaling branches and the  in planta  requirement of simultaneous phosphorylation at two key phosphorylation sites in SLAC1. We identify novel mechanisms ensuring specificity and robustness within stomatal Ca 2+ -signaling on a cellular, genetic, and biochemical level.

Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells