Eri Adams

Transport, signaling, and homeostasis of potassium and sodium in plants

Potassium (K⁺) is an essential macronutrient in plants and a lack of K⁺ significantly reduces the potential for plant growth and development. By contrast, sodium (Na⁺), while beneficial to some extent, at high concentrations it disturbs and inhibits various physiological processes and plant growth. Due to their chemical similarities, some functions of K⁺ can be undertaken by Na⁺ but K⁺ homeostasis is severely affected by salt stress, on the other hand. Recent advances have highlighted the fascinating regulatory mechanisms of K⁺ and Na⁺ transport and signaling in plants. This review summarizes three major topics: (i) the transport mechanisms of K⁺ and Na⁺ from the soil to the shoot and to the cellular compartments; (ii) the mechanisms through which plants sense and respond to K⁺ and Na⁺ availability; and (iii) the components involved in maintenance of K⁺/Na⁺ homeostasis in plants under salt stress.

Cesium Inhibits Plant Growth through Jasmonate Signaling in Arabidopsis thaliana

It has been suggested that cesium is absorbed from the soil through potassium uptake machineries in plants; however, not much is known about perception mechanism and downstream response. Here, we report that the jasmonate pathway is required in plant response to cesium. Jasmonate biosynthesis mutant aos and jasmonate-insensitive mutant coi1-16 show clear resistance to root growth inhibition caused by cesium. However, the potassium and cesium contents in these mutants are comparable to wild-type plants, indicating that jasmonate biosynthesis and signaling are not involved in cesium uptake, but involved in cesium perception. Cesium induces expression of a high-affinity potassium transporter gene HAK5 and reduces potassium content in the plant body, suggesting a competitive nature of potassium and cesium uptake in plants. It has also been found that cesium-induced HAK5 expression is antagonized by exogenous application of methyl-jasmonate. Taken together, it has been indicated that cesium inhibits plant growth via induction of the jasmonate pathway and likely modifies potassium uptake machineries.

Selective chemical binding enhances cesium tolerance in plants through inhibition of cesium uptake

High concentrations of cesium (Cs+) inhibit plant growth but the detailed mechanisms of Cs+ uptake, transport and response in plants are not well known. In order to identify small molecules with a capacity to enhance plant tolerance to Cs+, chemical library screening was performed using Arabidopsis. Of 10,000 chemicals tested, five compounds were confirmed as Cs+ tolerance enhancers. Further investigation and quantum mechanical modelling revealed that one of these compounds reduced Cs+ concentrations in plants and that the imidazole moiety of this compound bound specifically to Cs+. Analysis of the analogous compounds indicated that the structure of the identified compound is important for the effect to be conferred. Taken together, Cs+ tolerance enhancer isolated here renders plants tolerant to Cs+ by inhibiting Cs+ entry into roots via specific binding to the ion thus, for instance, providing a basis for phytostabilisation of radiocesium-contaminated farmland.

14-3-3 Proteins Participate in Light Signaling through Association with PHYTOCHROME INTERACTING FACTORs

14-3-3 proteins are regulatory proteins found in all eukaryotes and are known to selectively interact with phosphorylated proteins to regulate physiological processes. Through an affinity purification screening, many light-related proteins were recovered as 14-3-3 candidate binding partners. Yeast two-hybrid analysis revealed that the 14-3-3 kappa isoform (14-3-3κ) could bind to PHYTOCHROME INTERACTING FACTOR3 (PIF3) and CONSTITUTIVE PHOTOMORPHOGENIC1 (COP1). Further analysis by in vitro pull-down assay confirmed the interaction between 14-3-3κ and PIF3. Interruption of putative phosphorylation sites on the 14-3-3 binding motifs of PIF3 was not sufficient to inhibit 14-3-3κ from binding or to disturb nuclear localization of PIF3. It was also indicated that 14-3-3κ could bind to other members of the PIF family, such as PIF1 and PIF6, but not to LONG HYPOCOTYL IN FAR-RED1 (HFR1). 14-3-3 mutants, as well as the PIF3 overexpressor, displayed longer hypocotyls, and a pif3 mutant displayed shorter hypocotyls than the wild-type in red light, suggesting that 14-3-3 proteins are positive regulators of photomorphogenesis and function antagonistically with PIF3. Consequently, our results indicate that 14-3-3 proteins bind to PIFs and initiate photomorphogenesis in response to a light signal.

AtSKIP18 and AtSKIP31, F-box subunits of the SCF E3 ubiquitin ligase complex, mediate the degradation of 14-3-3 proteins in Arabidopsis

14-3-3 proteins regulate numerous cellular processes through interaction with their target proteins in a phosphorylation dependent manner. Although proteins that are regulated by 14-3-3s have been studied, the regulatory mechanism of 14-3-3s is poorly understood. In the present study, F-box proteins, a component of Skp1-Cullin-F-box E3 ubiquitin ligase, were identified as 14-3-3 targets using yeast two-hybrid screening. Among them, AtSKIP18 and AtSKIP31, were shown to mediate the degradation of Arabidopsis 14-3-3s. Mutational analyses of AtSKIP18 and AtSKIP31 indicated that the phosphorylation of AtSKIPs is critical for interaction and degradation of 14-3-3s. The loss-of-function mutation in AtSKIP31 resulted in enhanced primary root growth under nitrogen deficient conditions. These findings suggest that AtSKIP31 regulates the primary root growth in nitrogen deficiency via degrading 14-3-3s.

Overexpression of a Novel Component Induces HAK5 and Enhances Growth in Arabidopsis

Plants have developed mechanisms to adapt to the potassium deficient conditions over the years. In Arabidopsis thaliana, expression of a potassium transporter HAK5 is induced in low potassium conditions as an adaptive response to nutrient deficiency. In order to understand the mechanism in which HAK5 is regulated, the full-length cDNA overexpressor gene hunting system was employed as a screening method. Of 40 genes recovered, At4g18280 was found to be dramatically induced in response to potassium-deficiency and salt stress. Plants overexpressing this gene showed higher HAK5 expression and enhanced growth. These plants were also less sensitive to potassium-deficiency in terms of primary root growth. Taken together, these data suggest that this novel component, At4g18280, contributes to regulation of HAK5 and, consequently, tolerance to potassium-deficiency in plants.

Cesium Uptake in Plants: Mechanism, Regulation and Application for Phytoremediation

Radiocesium is mainly generated through anthropogenic activities and poses a great threat to health and the environment. As an alternative to costly physical and chemical methods to remediate the contaminated soil and water, phytoremediation, a technique making use of plants to remove or stabilise contamination, is receiving increasing attention. Selection of plant species that accumulate high levels of radiocesium has, therefore, been intensively investigated. In recent years, molecular techniques have enabled researchers to elucidate cesium transporters and regulatory mechanisms through which cesium uptake occurs in plants. Some proteins have been predicted to mediate cesium and many players, including cations and phytohormones, have been suggested as regulators. Although the molecular understanding of cesium uptake in plants is only just being revealed, the knowledge can readily be applied to radiocesium phytoremediation. In parallel, efforts to improve phytoremediation efficiency have sought the aid of biotic and abiotic factors such as microorganisms and chemicals. In this chapter, findings on the molecular mechanisms and regulation of cesium uptake in plants, what is known and what needs to be researched, are discussed. Then follows a discussion of the plant species suitable for radiocesium phytoremediation and the factors which improve phytoremediation efficiency.

A novel role for methyl cysteinate, a cysteine derivative, in cesium accumulation in Arabidopsis thaliana

Phytoaccumulation is a technique to extract metals from soil utilising ability of plants. Cesium is a valuable metal while radioactive isotopes of cesium can be hazardous. In order to establish a more efficient phytoaccumulation system, small molecules which promote plants to accumulate cesium were investigated. Through chemical library screening, 14 chemicals were isolated as ‘cesium accumulators’ in Arabidopsis thaliana. Of those, methyl cysteinate, a derivative of cysteine, was found to function within the plant to accumulate externally supplemented cesium. Moreover, metabolite profiling demonstrated that cesium treatment increased cysteine levels in Arabidopsis. The cesium accumulation effect was not observed for other cysteine derivatives or amino acids on the cysteine metabolic pathway tested. Our results suggest that methyl cysteinate, potentially metabolised from cysteine, binds with cesium on the surface of the roots or inside plant cells and improve phytoaccumulation.

Selection and functional analysis of a Pyropia yezoensis ammonium transporter PyAMT1 in potassium deficiency

Seaweeds are believed to have developed unique mechanisms to maintain optimal cellular potassium and sodium concentrations in order to survive in the saline marine environment. To gain a molecular understanding of underlying potassium/sodium homeostasis in seaweeds, full-length cDNA libraries from the multiple stages in the life cycle, including gametophytes, conchosporangia and sporophytes of a marine red alga, Pyropia yezoensis, were constructed. A large portion of genes from each library through the life cycle was revealed to be functionally unknown reconfirming the uniqueness of P. yezoensis genes in terms of evolutionary lineage. Genes that could potentially contribute to potassium deficiency tolerance were selected from the potassium uptake defective Escherichia coli strain expressing gametophytes and conchosporangia libraries under the low potassium conditions. Of those, an ammonium transporter gene, PyAMT1, was demonstrated to enhance potassium deficiency tolerance effectively when expressed in the E. coli strain. Potential roles of PyAMT1 and other candidate components in this context are discussed.