A transient outward-rectifying K(+ )channel current down-regulated by cytosolic Ca(2+ )in Arabidopsis thaliana guard cells

Sustained (noninactivating) outward-rectifying K(+) channel currents have been identified in a variety of plant cell types and species. Here, in Arabidopsis thaliana guard cells, in addition to these sustained K(+) currents, an inactivating outward-rectifying K(+) current was characterized (plant A-type current: I(AP)). I(AP) activated rapidly with a time constant of 165 ms and inactivated slowly with a time constant of 7.2 sec at +40 mV. I(AP) was enhanced by increasing the duration (from 0 to 20 sec) and degree (from +20 to −100 mV) of prepulse hyperpolarization. Ionic substitution and relaxation (tail) current recordings showed that outward I(AP) was mainly carried by K(+) ions. In contrast to the sustained outward-rectifying K(+) currents, cytosolic alkaline pH was found to inhibit I(AP) and extracellular K(+) was required for I(AP) activity. Furthermore, increasing cytosolic free Ca(2+) in the physiological range strongly inhibited I(AP) activity with a half inhibitory concentration of ≈ 94 nM. We present a detailed characterization of an inactivating K(+) current in a higher plant cell. Regulation of I(AP) by diverse factors including membrane potential, cytosolic Ca(2+) and pH, and extracellular K(+) and Ca(2+) implies that the inactivating I(AP) described here may have important functions during transient depolarizations found in guard cells, and in integrated signal transduction processes during stomatal movements.