Regulation of leaf hydraulics: from molecular to whole plant levels

The water status of plant leaves is dependent on both stomatal regulation and water supply from the vasculature to inner tissues. The present review addresses the multiple physiological and mechanistic facets of the latter process. Inner leaf tissues contribute to at least a third of the whole resistance to water flow within the plant. Physiological studies indicated that leaf hydraulic conductance (K-leaf) is highly dependent on the anatomy, development and age of the leaf and can vary rapidly in response to physiological or environmental factors such as leaf hydration, light, temperature, or nutrient supply. Differences in venation pattern provide a basis for variations in Kleaf during development and between species. On a short time (hour) scale, the hydraulic resistance of the vessels can be influenced by transpiration-induced cavitations, wall collapses, and changes in xylem sap composition. The extravascular compartment includes all living tissues (xylem parenchyma, bundle sheath, and mesophyll) that transport water from xylem vessels to substomatal chambers. Pharmacological inhibition and reverse genetics studies have shown that this compartment involves water channel proteins called aquaporins (AQPs) that facilitate water transport across cell membranes. In many plant species, AQPs are present in all leaf tissues with a preferential expression in the vascular bundles. The various mechanisms that allow adjustment of Kleaf to specific environmental conditions include transcriptional regulation of AQPs and changes in their abundance, trafficking, and intrinsic activity. Finally, the hydraulics of inner leaf tissues can have a strong impact on the dynamic responses of leaf water potential and stomata, and as a consequence on plant carbon economy and leaf expansion growth. The manipulation of these functions could help optimize the entire plant performance and its adaptation to extreme conditions over short and long time scales.