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Stomata begin to close within 24 h of imposing soil flooding. We investigated whether the stomatal response was triggered by reduced photosynthetic efficiency in young, fully expanded leaves of flooded plants. Chlorophyll fluorescence measurements indicated that ФpsII the effective quantum yield of PS II,decreased after stomata began to close in flooded plants. Changes in qP mirrored those of ФpsII, ФpsII was not affected by daytime patterns of stomatal conductance in well-drained plants but was reduced by stomatal closure in flooded plants. Fv/Fm a measure of the overall photosynthetic efficiency of dark- adapted plants, decreased after 57 h of flooding. Therefore, prolonged soil flooding adversely affected the thylakoid membranes. QN, a measure of the amount of captured energy dissipated as heat and therefore, unused by the photosynthetic machinery, began to increase after 32 h of flooding and continued to rise thereafter. The interdependence of the changes in chlorophyll fluorescence parameters and the flooding-induced closure of stomata is discussed.
Flooding of the soil induces stomata to close within a few hours decreasing a potential damage of leaves which would otherwise occur because of a decrease of root hydraulic conductivity. The signals triggering these shoot responses have not been fully identified but could include changes in hormone transport from roots to shoots as their synthesis and xylem loading are altered. The present research determined if changes in the delivery of indole acetic acid (IAA) could be a root-born signal comparable with decreases in abscisic acid (ABA) reported previously. Tomato plants at the 7-8-leaf stage were flooded up to 48 h by submerging their pots individually in tap water. Xylem sap was collected from freshly detopped and pneumatically pressurised roots at flow rates equivalent to those of whole-plant transpiration. Concentrations of ABA and IAA in the sap were quantified by the GC-MS and their delivery rates from roots to shoots (fluxes) was calculated an the basis of transpiration rates of the whole plant calculating the amounts delivered per unit area of leaf. Leaf conductance (a measure of stomatal closure) and leaf water potential (LWP) were also measured. Stomata were closed almost completely in 8 h of flooding. This decreased transpiration significantly. Stomatal closure and transpiration remained much below than those of well-drained plants for at least 48 h. Between 4-8 h of flooding, a marked transient decrease in LWP took place which was quickly succeeded by its increase even to values above those of well-drained plants. The concentration and the delivery of ABA from flooded roots to shoots in xylem sap decreased 5- and 7-fold, respectively, within 2 h of flooding and remained lower for at least 48 h, as compared to the control. In contrast, IAA concentrations in xylem sap of flooded plants were above those of well-drained plants. However, slower rates of transpiration generated IAA deliveries that were decreased by 36, 35, 18 and 28% after 2, 4, 6 and 8 h of flooding respectively. The rapid and transient decrease in LWP of leaves and decrease in the delivery of ABA and IAA from roots to shoots during the first hours of flooding were always observed before the stomata started to close. Each is a potentially active signal. While the decrease in ABA is diffucult to explain the stomatal closure, the involvement of the rapidly depressed delivery rates of IAA from roots to shoots during the early flooding merits further investigation.
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