Surrounded by luxury: The necessities of subsidiary cells.

The evolution of stomata marks one of the key advances that enabled plants to colonise dry land while allowing gas exchange for photosynthesis. In large measure, stomata retain a common design across species that incorporates paired guard cells with little variation in structure. By contrast, the cells of the stomatal complex immediately surrounding the guard cells vary widely in shape, size and count. Their origins in development are similarly diverse. Thus, the surrounding cells are likely a luxury that the necessity of stomatal control cannot do without (with apologies to Oscar Wilde). Surrounding cells are thought to support stomatal movements as solute reservoirs and to shape stomatal kinetics through backpressure on the guard cells. Their variety may also reflect a substantial diversity in function. Certainly modelling, kinetic analysis and the few electrophysiological studies to date give hints of much more complex contributions in stomatal physiology. Even so, our knowledge of the cells surrounding the guard cells in the stomatal complex is far from complete.

Engineering stomata for enhanced carbon capture and water-use efficiency.

Stomatal pores facilitate gaseous exchange between the inner air spaces of the leaf and the atmosphere. As gatekeepers that balance CO2 entry for photosynthesis against transpirational water loss, they are a focal point for efforts to improve crop performance, especially in the efficiency of water use, within the changing global environment. Until recently, engineering strategies had focused on stomatal conductance in the steady state. These strategies are limited by the physical constraints of CO2 and water exchange such that gains in water-use efficiency (WUE) commonly come at a cost in carbon assimilation. Attention to stomatal speed and responsiveness circumvents these constraints and offers alternatives to enhancing WUE that also promise increases in carbon assimilation in the field.

OnGuard3e: A predictive, ecophysiology-ready tool for gas exchange and photosynthesis research.

Gas exchange across the stomatal pores of leaves is a focal point in studies of plant-environmental relations. Stomata regulate atmospheric exchange with the inner air spaces of the leaf. They open to allow CO2 entry for photosynthesis and close to minimize water loss. Models that focus on the phenomenology of stomatal conductance generally omit the mechanics of the guard cells that regulate the pore aperture. The OnGuard platform fills this gap and offers a truly mechanistic approach with which to analyse stomatal gas exchange, whole-plant carbon assimilation and water-use efficiency. Previously, OnGuard required specialist knowledge of membrane transport, signalling and metabolism. Here we introduce OnGuard3e, a software package accessible to ecophysiologists and membrane biologists alike. We provide a brief guide to its use and illustrate how the package can be applied to explore and analyse stomatal conductance, assimilation and water use efficiencies, addressing a range of experimental questions with truly predictive outputs.

The outward shaker channel OsK5.2 improves plant salt tolerance by contributing to control of both leaf transpiration and K+ secretion into xylem sap.

Soil salinity constitutes a major environmental constraint to crop production worldwide. Leaf K+ /Na+ homoeostasis, which involves regulation of transpiration, and thus of the xylem sap flow, and control of the ionic composition of the ascending sap, is a key determinant of plant salt tolerance. Here, we show, using a reverse genetics approach, that the outwardly rectifying K+ -selective channel OsK5.2, which is involved in both K+ release from guard cells for stomatal closure in leaves and K+ secretion into the xylem sap in roots, is a strong determinant of rice salt tolerance (plant biomass production and shoot phenotype under saline constraint). OsK5.2 expression was upregulated in shoots from the onset of the saline treatment, and OsK5.2 activity in guard cells led to a fast decrease in transpirational water flow and, therefore, reduced Na+ translocation to shoots. In roots, upon saline treatment, OsK5.2 activity in xylem sap K+ loading was maintained, and even transiently increased, outperforming the negative effect on K+ translocation to shoots resulting from the reduction in xylem sap flow. Thus, the overall activity of OsK5.2 in shoots and roots, which both reduces Na+ translocation to shoots and benefits shoot K+ nutrition, strongly contributes to leaf K+ /Na+ homoeostasis.

A Dual Role for the OsK5.2 Ion Channel in Stomatal Movements and K+ Loading into Xylem Sap.

The roles of potassium channels from the Shaker family in stomatal movements have been investigated by reverse genetics analyses in Arabidopsis (Arabidopsis thaliana), but corresponding information is lacking outside this model species. Rice (Oryza sativa) and other cereals possess stomata that are more complex than those of Arabidopsis. We examined the role of the outward Shaker K+ channel gene OsK5.2. Expression of the OsK5.2 gene (GUS reporter strategy) was observed in the whole stomatal complex (guard cells and subsidiary cells), root vasculature, and root cortex. In stomata, loss of OsK5.2 functional expression resulted in lack of time-dependent outward potassium currents in guard cells, higher rates of water loss through transpiration, and severe slowdown of stomatal closure. In line with the expression of OsK5.2 in the plant vasculature, mutant plants displayed a reduced K+ translocation from the root system toward the leaves via the xylem. The comparison between rice and Arabidopsis show that despite the strong conservation of Shaker family in plants, substantial differences can exist between the physiological roles of seemingly orthologous genes, as xylem loading depends on SKOR and stomatal closure on GORK in Arabidopsis, whereas both functions are executed by the single OsK5.2 Shaker in rice.

Differential expression analysis of a subset of GmNAC genes in shoots of two contrasting drought-responsive soybean cultivars DT51 and MTD720 under normal and drought conditions.

NAC transcription factors are known to be involved in regulation of plant responses to drought stress. In this study, the expression of 23 drought-responsive GmNAC genes was assessed in the shoot tissues of DT51 and MTD720, the two soybean varieties with contrasting drought-responsive phenotypes, by real-time quantitative PCR (RT-qPCR) under normal and drought conditions. Results indicated that expression profile of GmNAC genes was genotype-dependent, and six GmNACs (GmNAC019, 043, 062, 085, 095 and 101) had higher transcript levels in the shoots of the drought-tolerant DT51 in comparison with the drought-sensitive MTD720 under drought. Our study suggests a positive correlation between the higher drought tolerance degree of DT51 versus MTD720 and the up-regulation of at least these six drought-responsive GmNACs in the shoot tissues. Furthermore, on the basis of our analysis, three genes, GmNAC043, 085 and 101, were identified as promising candidates for development of drought-tolerant soybean cultivars by genetic engineering.