Inventory and functional characterization of the HAK potassium transporters of rice

Plants take up large amounts of K+ from the soil solution and distribute it to the cells of all organs, where it fulfills important physiological functions. Transport of K+ from the soil solution to its final destination is mediated by channels and transporters. To better understand K+ movements in plants, we intended to characterize the function of the large KT-HAK-KUP family of transporters in rice (Oryza saliva cv Nipponbare). By searching in databases and cDNA cloning, we have identified 17 genes (OsHAK1-17) encoding transporters of this family and obtained evidence of the existence of other two genes. Phylogenetic analysis of the encoded transporters reveals a great diversity among them, and three distant transporters, OSHAK1, OsHAK7, and OsHAK10, were expressed in yeast (Saccharomyces cerevisiae) and bacterial mutants to determine their functions. The three transporters mediate K+ influxes or effluxes, depending on the conditions of the experiment. A comparative kinetic analysis of HAK-mediated K+ influx in yeast and in roots of K+-starved rice seedlings demonstrated the involvement of HAK transporters in root K+ uptake. We discuss that all HAK transporters may mediate K+ transport, but probably not only in the plasma membrane. Transient expression of the OSHAK10-green fluorescent protein fusion protein in living onion epidermal cells targeted this protein to the tonoplast.