Potassium is an essential mineral element for plant growth and development. Although it is known that plants absorb and transport K+ through membrane transporters, it remains unclear how these transporters are regulated. Here we show that the protein kinase CIPK23, encoded by the LKS1 gene, regulates K+ uptake under low-K+ conditions. Lesion of LKS1 significantly reduced K+ uptake and caused leaf chlorosis and growth inhibition, whereas overexpression of LKS1 significantly enhanced K+ uptake and tolerance to low K+. We demonstrate that CIPK23 directly phosphorylates the K+ transporter AKT1 and further find that CIPK23 is activated by the binding of two calcineurin B-like proteins, CBL1 and CBL9. We propose a model in which the CBL1/9-CIPK23 pathway ensures activation of AKT1 and enhanced K+ uptake under low-K+ conditions.

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Potassium transporters play crucial roles in K + uptake and translocation in plants. However, so far little is known about the regulatory mechanism of potassium transporters. Here, we show that a Shaker-like potassium channel AtKC1, encoded by the AtLKT1 gene cloned from the Arabidopsis thaliana low-K + (LK)-tolerant mutant Atlkt1, significantly regulates AKT1-mediated K + uptake under LK conditions. Under LK conditions, the Atkc1 mutants maintained their root growth, whereas wild-type plants stopped their root growth. Lesion of AtKC1 significantly enhanced the tolerance of the Atkc1 mutants to LK stress and markedly increased K + uptake and K + accumulation in the Atkc1-mutant roots under LK conditions. Electrophysiological results showed that AtKC1 inhibited the AKT1-mediated inward K + currents and negatively shifted the voltage dependence of AKT1 channels. These results demonstrate that the 'silent' K + channel α-subunit AtKC1 negatively regulates the AKT1-mediated K + uptake in Arabidopsis roots and consequently alters the ratio of root-to-shoot under LK stress conditions. Keywords: Arabidopsis; potassium channel; low-K + stress; AKT1; AtKC1 Cell Research (2010) IntroductionPotassium is an essential mineral element for plant growth and development, and it plays essential roles in many important physiological and biochemical processes in living plant cells, such as regulation of enzyme activation, electrical neutralization, osmoregulation, control of membrane potential, co-transport of sugars, and so on [1,2]. Plant growth and development need millimolar K + in the soil or growth medium, but typical K + concentration at the interface of roots and soils is within micromolar range [3]. Thus, plants often encounter low-K + (LK) stress under natural conditions. Although different plants or different genotypes of a plant species show varied K + utilization efficiency [4], most plants show K + -deficient symptom under LK stress, typically leaf chlorosis and subsequent inhibition of plant growth and development [5].Absorption of K + by plant cells and K + translocation between different tissues and organs in plants are mediated by plant K + transporters and channels [2,6,7]. Over the past decade, a large number of genes encoding plant K + transporters and channels, particularly for Arabidopsis, have been characterized [6][7][8]. These K + transporters vary in K + affinity, kinetics, transcriptional modulation, regulatory mechanism, etc [2,[6][7][8], and they compose a complex system for plant K + uptake and translocation. Among these K + transporters and channels, members of the Shaker K + channel family are well characterized for their potential functions and are probably the most important for K + uptake and transport in Arabidopsis [8] Recent reports showed that channel heterotetramerization is a key regulatory mechanism for K + channels, which was found not only in animals [14][15][16] but also in plants [17,18]. Different kinds of potassium channel subunits may assemble together to form functional heteromer...