The family of plant membrane transporters named HKT (for high-affinity K+ transporters) can be subdivided into subfamilies 1 and 2, which, respectively, comprise Na+-selective transporters and transporters able to function as Na+-K+ symporters, at least when expressed in yeast (Saccharomyces cerevisiae) or Xenopus oocytes. Surprisingly, a subfamily 2 member from rice (Oryza sativa), OsHKT2;4, has been proposed to form cation/K+ channels or transporters permeable to Ca2+ when expressed in Xenopus oocytes. Here, OsHKT2;4 functional properties were reassessed in Xenopus oocytes. A Ca2+ permeability through OsHKT2;4 was not detected, even at very low external K+ concentration, as shown by highly negative OsHKT2;4 zero-current potential in high Ca2+ conditions and lack of sensitivity of OsHKT2;4 zero-current potential and conductance to external Ca2+. The Ca2+ permeability previously attributed to OsHKT2;4 probably resulted from activation of an endogenous oocyte conductance. OsHKT2;4 displayed a high permeability to K+ compared with that to Na+ (permeability sequence: K+ > Rb+ ≈ Cs+ > Na+ ≈ Li+ ≈ NH4  +). Examination of OsHKT2;4 current sensitivity to external pH suggested that H+ is not significantly permeant through OsHKT2;4 in most physiological ionic conditions. Further analyses in media containing both Na+ and K+ indicated that OsHKT2;4 functions as K+-selective transporter at low external Na+, but transports also Na+ at high (>10 mm) Na+ concentrations. These data identify OsHKT2;4 as a new functional type in the K+ and Na+-permeable HKT transporter subfamily. Furthermore, the high permeability to K+ in OsHKT2;4 supports the hypothesis that this system is dedicated to K+ transport in the plant.

Summary In grapevine, climate changes lead to increased berry potassium (K+) contents that result in must with low acidity. Consequently, wines are becoming ‘flat’ to the taste, with poor organoleptic properties and low potential aging, resulting in significant economic loss. Precise investigation into the molecular determinants controlling berry K+ accumulation during its development are only now emerging. Here, we report functional characterization by electrophysiology of a new grapevine Shaker‐type K+ channel, VvK3.1. The analysis of VvK3.1 expression patterns was performed by qPCR and in situ hybridization. We found that VvK3.1 belongs to the AKT2 channel phylogenetic branch and is a weakly rectifying channel, mediating both inward and outward K+ currents. We showed that VvK3.1 is highly expressed in the phloem and in a unique structure located at the two ends of the petiole, identified as a pulvinus. From the onset of fruit ripening, all data support the role of the VvK3.1 channel in the massive K+ fluxes from the phloem cell cytosol to the berry apoplast during berry K+ loading. Moreover, the high amount of VvK3.1 transcripts detected in the pulvinus strongly suggests a role for this Shaker in the swelling and shrinking of motor cells involved in paraheliotropic leaf movements.