Stomatal guard cells are widely recognized as the premier plant cell model for membrane transport, signaling, and homeostasis. This recognition is rooted in half a century of research into ion transport across the plasma and vacuolar membranes of guard cells that drive stomatal movements and the signaling mechanisms that regulate them. Stomatal guard cells surround pores in the epidermis of plant leaves, controlling the aperture of the pore to balance CO entry into the leaf for photosynthesis with water loss via transpiration. The position of guard cells in the epidermis is ideally suited for cellular and subcellular research, and their sensitivity to endogenous signals and environmental stimuli makes them a primary target for physiological studies. Stomata underpin the challenges of water availability and crop production that are expected to unfold over the next 20 to 30 years. A quantitative understanding of how ion transport is integrated and controlled is key to meeting these challenges and to engineering guard cells for improved water use efficiency and agricultural yields.

The major plant nutrient magnesium (Mg) is involved in numerous physiological processes and its deficiency can severely reduce the yield and quality of crops. Since Mg availability in soil and uptake into the plant is often limited by unfavorable soil or climatic conditions, application of Mg onto leaves, the site with highest physiological Mg demand, might be a reasonable alternative fertilization strategy. This study aimed to investigate, if MgSO4 leaf-application in practically relevant amounts can efficiently alleviate the effects of Mg starvation in maize, namely reduced photosynthesis capacity, disturbed ion homeostasis and growth depression. Results clearly demonstrated that Mg deficiency could be mitigated by MgSO4 leaf-application as efficiently as by resupply of MgSO4 via the roots in vegetative maize plants. Significant increases in SPAD values and net rate of CO2-assimilation as well as enhanced shoot biomass have been achieved. Ion analysis furthermore revealed an improvement of the nutrient status of Mg-deficient plants with regard to [Mg], [K], and [Mn] in distinct organs, thereby reducing the risk of Mn-toxicity at the rootside, which often occurs together with Mg deficiency on acid soils. In conclusion, foliar fertilization with Mg proved to be an efficient strategy to adequately supply maize plants with Mg and might hence be of practical relevance to correct nutrient deficiencies during the growing season.

Stomata of most plants close to preserve water when the demand for CO2 by photosynthesis is reduced. Stomatal responses are slow compared to photosynthesis, and this kinetic difference erodes assimilation and water use efficiencies under fluctuating light. Despite a deep knowledge of guard cells that regulate the stoma, efforts to enhance stomatal kinetics are limited by our understanding of its control by foliar CO2. Guided by mechanistic modelling that incorporates foliar CO2 diffusion and mesophyll photosynthesis, here we uncover a central role for endomembrane Ca 2+ stores in guard cell responsiveness to fluctuating light and CO2. Modelling predicted and experiments demonstrated a delay in Ca 2+cycling that was enhanced by endomembrane Ca 2+ -ATPase mutants, altering stomatal conductance and reducing assimilation and water use efficiency. Our findings illustrate the power of modelling to bridge the gap from the guard cell to whole-plant photosynthesis, and they demonstrate an unforeseen latency, or 'carbon memory', of guard cells that affects stomatal dynamics, photosynthesis and water use efficiency.

Models of guard cell dynamics, built on the OnGuard platform, have provided quantitative insights into stomatal function, demonstrating substantial predictive power. However, the kinetics of stomatal opening predicted by OnGuard models were threefold to fivefold slower than observed in vivo. No manipulations of parameters within physiological ranges yielded model kinetics substantially closer to these data, thus highlighting a missing component in model construction. One well‐documented process influencing stomata is the constraining effect of the surrounding epidermal cells on guard cell volume and stomatal aperture. Here, we introduce a mechanism to describe this effect in OnGuard2 constructed around solute release and a decline in turgor of the surrounding cells and its subsequent recovery during stomatal opening. The results show that this constraint–relaxation–recovery mechanism in OnGuard2 yields dynamics that are consistent with experimental observations in wild‐type Arabidopsis, and it predicts the altered opening kinetics of ost2 H+‐ATPase and slac1 Cl− channel mutants. Thus, incorporating solute flux of the surrounding cells implicitly through their constraint on guard cell expansion provides a satisfactory representation of stomatal kinetics, and it predicts a substantial and dynamic role for solute flux across the apoplastic space between the guard cells and surrounding cells in accelerating stomatal kinetics.

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