Leaf hydraulic maze; Differential effect of ABA on bundle-sheath, palisade, and spongy mesophyll hydraulic conductance

Abstract

ABSTRACTLeaf hydraulic conductance (Kleaf) facilitates the movement of water for transpiration, enabling continual CO2uptake while the plant maintains its water status. We hypothesized that bundle-sheath and mesophyll cells play key roles in regulating the radial flow of water out of the xylem under optimal and stress conditions. To examine that hypothesis, we generated transgenicArabidopsisplants that were insensitive to abscisic acid (ABA) in their bundle sheath (BSabi) or mesophyll (MCabi) cells. Both BSabi and MCabi plants showed greater Kleafand transpiration under optimal conditions. Yet, the stomatal apertures, stomatal indices and vein densities of the BSabi plants were similar to those of WT plants. MCabi plants had larger stomatal apertures, a higher stomatal index and greater vascular diameter and biomass, relative to the WT and BSabi. In response to xylem-fed ABA, both transgenic and WT plants reduced their Kleafand transpiration. However, leaf water potential was reduced only in the WT. The membrane osmotic water permeability (Pf) of the WTs’ spongy mesophyll was higher than that of its palisade mesophyll. Moreover, only the spongy cells reduced theirPfin response to ABA. ABA-insensitive spongy mesophyll cells had a lowPf; whereas ABA-insensitive bundle-sheath cells had a higherPf.Palisade cells maintained a lowPfat all ABA levels. ABA increased the symplastic water pathway, but its contribution to Kleafwas negligible. We suggest that the bundle sheath–spongy mesophyll pathway may control Kleafto maintain steady-state conditions in the palisade cells and optimal whole-leaf water-use efficiency.