Stomatal closure under drought is controlled by below-ground hydraulics 

Abstract

<p>Stomatal closure allows plants to promptly respond to water shortage. Although the coordination between stomatal regulation, leaf and xylem hydraulics has been extensively investigated, the impact of below-ground hydraulics on stomatal regulation remains unknown.</p><p>We used a novel root pressure chamber to measure, during soil drying, the relation between transpiration rate (<em>E</em>) and leaf xylem water pressure (<em>ψ</em><sub>leaf-x</sub>) in tomato shoots grafted onto two contrasting rootstocks, a long and a short one. In parallel, we also measured the <em>E</em>(<em>ψ</em><sub>leaf-x</sub>) relation without pressurization. A soil–plant hydraulic model was used to reproduce the measurements. We hypothesize that (1) stomata close when the <em>E</em>(<em>ψ</em><sub>leaf-x</sub>) relation becomes non-linear and (2) non-linearity occurs at higher soil water contents and lower transpiration rates in short-rooted plants.</p><p>The <em>E</em>(<em>ψ</em><sub>leaf-x</sub>) relation was linear in wet conditions and became non-linear as the soil dried. Changing below-ground traits (i.e. root system) significantly affected the <em>E</em>(<em>ψ</em><sub>leaf-x</sub>) relation during soil drying. Plants with shorter root systems required larger gradients in soil water pressure to sustain the same transpiration rate and exhibited an earlier non-linearity and stomatal closure.</p><p>We conclude that, during soil drying, stomatal regulation is controlled by below-ground hydraulics in a predictable way. The model suggests that the loss of hydraulic conductivity occurred in soil. These results prove that stomatal regulation is intimately tied to root and soil hydraulic conductances.</p>