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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.
Hydrogen peroxide (H2O2) is produced predominantly in plant cells during photosynthesis and photorespiration, and to a lesser extent, in respiration processes. It is the most stable of the so– called reactive oxygen species (ROS), and therefore plays a crucial role as a signalling molecule in various physiological processes. Intra- and intercellular levels of H2O2 increase during environmental stresses. Hydrogen peroxide interacts with thiol-containing proteins and activates different signalling pathways as well as transcription factors, which in turn regulate gene expression and cell-cycle processes. Genetic systems controlling cellular redox homeostasis and H2O2 signalling are discussed. In addition to photosynthetic and respiratory metabolism, the extracellular matrix (ECM) plays an important role in the generation of H2O2, which regulates plant growth, development, acclimatory and defence responses. During various environmental stresses the highest levels of H2O2 are observed in the leaf veins. Most of our knowledge about H2O2 in plants has been obtained from obligate C3 plants. The potential role of H2O2 in the photosynthetic mode of carbon assimilation, such as C4 metabolism and CAM (Crassulacean acid metabolism) is discussed. We speculate that early in the evolution of oxygenic photosynthesis on Earth, H2O2 could have been involved in the evolution of modern photosystem II.
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