Structure and function of potassium channels in plants: some inferences about the molecular origin of inward rectification in KAT1 channels (Review)

Potassium channels in plants play a variety of important physiological roles including K(+) uptake into roots, stomatal and leaf movements, and release of K(+) into the xylem. This review summarizes current knowledge about a class of plant genes whose products are K(+) channel-forming proteins. Potassium channels of this class belong to a superfamily characterized by six membrane-spanning domains (S1-6), a positively charged S4 domain and a region between the S5 and S6 segments that forms the channel selectivity filter. These channels are voltage dependent, which means the membrane potential modifies the probability of opening (P(o)). However, despite these channels sharing the same topology as the outward-rectifying K(+) channels, which are activated by membrane depolarization, some plant K(+) channels such as KAT1/2 and KST1 open with hyperpolarizing voltages. In outward-rectifying K(+) channels, the change in P(o) is achieved through a voltage sensor formed by the S4 segment that detects the voltage transferring its energy to the gate that controls pore opening. This coupling is achieved by an outward displacement of the charges contained in S4. In KAT1, most of the results indicate that S4 is the voltage sensor. However, how the movement of S4 leads to opening remains unanswered. On the basis of recent data, we propose here that in plant-inward rectifiers an inward movement of S4 leads to channel opening and that the difference between it and outward-rectifying channels resides in the mechanism that couples gating charge displacement with pore opening.