When the night is not long enough: soil drying constrains nighttime tree rehydration, setting a physical limit to stomatal regulation

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Abstract
Plants are often assumed to optimize carbon assimilation against water loss through stomatal regulation, responding instantaneously to varying atmospheric demand and soil water supply. However, a growing body of work highlights the importance of nighttime recovery processes for understanding plant water use, yet nighttime processes are not considered in existing supply–demand models. Here, we extend an existing supply–demand model by including flux imbalances between atmospheric demand and soil supply buffered by plant water storage. The flux imbalances accumulate during the day and are compensated overnight. Applying the conceptual model to field data from Quercus pubescens Willd., we show that progressive soil drying constrains the rate of nighttime rehydration. Precisely, below a critical soil water potential threshold (ψsoil,crit), nighttime rehydration begins to decline because rehydration rates become too low relative to the duration of the night (‘the night is not long enough’). This threshold marks a transition beyond which maintaining high transpiration rates becomes increasingly risky. The observations show that stomatal regulation begins near this threshold, pointing to the concomitance between daytime stomatal regulation and limited nighttime rehydration. These results suggest that progressive soil drying constrains nighttime rehydration and reveal a critical hydraulic threshold that coincides with the onset of stomatal regulation and highlight that plant water use is constrained not only by daytime fluxes but also by nighttime recovery.

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