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This work reports on experimental evidence for the role of ion-mediated changes of xylem hydraulic conductivity in the functional response of Solanum lycopersicum L. cv. Naomi to moderate salinity levels. Measurements were performed in fully developed 12-week-old plants grown in half-strength Hoagland solution (control, C-plants) or in the same solution added with 35mM NaCl (NaCl-plants). NaCl-plants produced a significantly less but heavier leaves and fruits but had similar gas-exchange rates as control plants. Moreover, NaCl-plants showed higher vessel multiple fraction (FVM) than control plants. Xylem sap potassium and sodium concentrations were significantly higher in NaCl-plants than in control plants. When stems were perfused with 10mM NaCl or KCl, the hydraulic conductance of NaCl plants was nearly 1.5 times higher than in control plants. Accordingly, stem hydraulic conductance measured in planta was higher in NaCl- than in control plants. Our data suggest that tomato plants grown under moderate salinity upregulate xylem sap [Na+] and [K+], as well as sensitivity of xylem hydraulics to sap ionic content, thus, increasing water transport capacity.

Hydraulic conductance (Kleaf) and morpho-anatomical parameters were measured in sun and shade Quercus ilex L. (holm oak) leaves. Sun leaves had lower surface area (Aleaf) and volume (Vleaf) and higher specific mass (leaf mass per area, LMA) than shade leaves. Transpiration rate and Kleaf scaled by Aleaf (Kleaf_area) were 2-fold higher in sun than in shade leaves. Kleaf_area was not correlated with vein density or stomatal density, which were found to be similar in the two leaf types. Values of Kleaf scaled by Vleaf or leaf dry weight (Kleaf_dw) were only 40% higher in sun than in shade leaves, suggesting that structural changes of Holm oak leaves acclimating to different light intensities enhance water transport to the unit evaporating leaf surface area, while maintaining more constant hydraulic supply to mesophyll cells and carbon costs of the water transport system. Sun leaves had higher Kleaf_dw and LMA than shade ones, indicating that high LMA resulted from resource allocation involved in both water transport and structural rigidity. Future studies of the intra- and inter-specific variability of mass-based hydraulic efficiency might provide important insights into leaf hydraulics and carbon economy. Kleaf_dw might prove to be an important driver of plant acclimation and adaptation to the environment.

Xylem cavitation is a common occurrence in drought-stressed plants. Cavitation-induced embolism reduces xylem hydraulic conductivity (kxylem) and may lead to stomatal closure and reduction of photosynthetic rates. Recent studies have suggested that plants may compensate for kxylem loss through ion-mediated enhancement of the residual water transport capacity of functioning conduits. To test this hypothesis, field-grown laurel (Laurus nobilis L.) plants were subjected to mild drought stress by suspending irrigation. Drought treatment induced a significant increase in xylem embolism compared with control (well watered) plants. Xylem sap potassium concentration ([K+]) increased during the day both in control and water stressed plants. Midday values of sap [K+] were significantly higher in water stressed plants. The recorded increase in sap potassium concentration induced significant enhancement of residual kxylem when solutions with different [K+] were perfused through excised stems sampled in the field and measured in the laboratory. In planta measurements of stem hydraulic conductance revealed no change between water stressed plants and controls. Our data suggest that ion-mediated enhancement of residual kxylem buffered the actual loss of hydraulic conductance suffered by plants during the warmest hours of the day as well as under mild drought stress conditions.

The ability of stems of Laurus nobilis (L.) to refill embolised xylem conduits was studied in plants both at optimal water supply (W) and under conditions of soil drought inducing xylem pressures (P x ) of -1.54 (S1) and -2.35 MPa (S2). Starch depolymerisation in wood parenchyma was measured as percentage of cells 'with high starch content' (HSC-cells) counted under a microscope. HSC-cells decreased during embolism and increased again in refilled stems. A direct relationship was found between percentage of HSC-cells and P x , with HSC-cells between 65 and 75% of the total at P x  ≥ -0.6 MPa, at which recovery from PLC was recorded. At low transpiration, starch re-appeared in wood parenchyma cells but only in plants that showed diurnal stomatal opening (W- and S1-plants). In S2-plants showing diurnal stomatal closure and nocturnal opening with P x between -1.2 to -2.4 MPa, HSC-cells were only 25% and plants did not recover from PLC. This finding suggests that (i) the P x threshold for embolism repair was ≥ -0.6 MPa, and (ii) impeded phloem loading limits starch content in wood parenchyma and embolism repair. We conclude that starch depolymerisation acts as a signal to phloem unloading sugars to embolised conduits thus generating the necessary osmotic gradients driving refilling.

Xylem embolism repair is preceded by starch depolymerisation in vessel-associated cells (VAC) of Laurus nobilis L. (laurel) twigs, but the primary signal triggering such a process is still unknown. We tested the hypothesis that conduit wall vibrations during cavitation may be sensed by VAC inducing starch-to-sugar conversion. Twigs of laurel from watered or stressed plants were exposed to ultrasound for 60 min to simulate acoustic waves emitted by cavitating conduits. Preliminary tests showed that ultrasound caused no damage to cell membrane integrity nor did they cause xylem embolism. The number of VAC with high starch content (HSC-cells) was estimated microscopically by counting the cells with more than 50% of their lumen filled with starch granules. Sonication had no effect on HSC-cells in twigs from watered plants while it induced a drop in the percentage HSC-cells from 80 to 40% in twigs from stressed plants, at the ultrasound source location. No effect was recorded in these twigs 20 mm from the ultrasound source. Sonication was a good simulator of cavitation in inducing starch depolymerisation which suggests a possible bio- physical nature for the signal initiating embolism repair.

Phoma trachephila (Petri) Kantschaveli et Gikachvili causes dieback of several Citrus species. The impact of this fungus on leaf hydraulics was studied in Citrus aurantium L. (sour orange) with the aim of identifying the primary mechanism of damage to leaves. Leaves inoculated with a conidial suspension were measured for conductance to water vapor (g L ) and specific hydraulic conductance (K leaf ) every 3 days after inoculation. The earliest symptom of infection consisted of vein chlorosis. Functional vein density (FVD) was monitored and microscopic observations were made of major vein conduits. Impairment of vein hydraulics started 25 days after inoculation with a losses of K leaf of 40% and g L of ~60%. Most minor veins within chlorotic areas were no longer functioning and some conduits of the major veins showed digested interconduit pits leading to vein cavitation. The close K leaf -FVD relationship revealed that vein impairment caused drop of K leaf and, consequently, of g L at chlorotic areas. Leaf infection was focused to veins that were first forced to embolise and then invaded by fungal hyphae. The vein embolism due to the Phoma amplifies the native dominant hydraulic resistance of leaf veins, and leads ultimately to early shedding of infected leaves.

Xylem recovery from embolism was studied in stems of Laurus nobilis L. that were induced to cavitate by combining negative xylem pressures with positive air pressures applied with a pressure collar. Xylem refilling was measured 2 and 20 min and 15 h after air pressure release in January, March and June when increasing percentages of wood parenchyma cells with high starch content (HSC-VAC) were counted (from 0% in January to 87.3% in June). In January, no xylem repair was measured. In June, stems refilled by 75% of previous conductivity loss with a parallel decrease of HSC-VAC. Xylem refilling was tested for stems with phloem either intact or excised by 20 and 50% and with phloem inactivated by girdling stems at both sides of the embolised segment. Stems with 50% of the cortex removed showed some recovery 15 h after embolism. Girdled stems did not recover from embolism and no starch depolymerisation was measured. Girdled stems where a radial mechanical pressure was applied for 20 min after embolism refilled in the same way as stems with intact phloem. Our conclusion is that phloem may export some signal for starch depolymerisation and this, in turn, would drive sugar efflux into embolised conduits with consequent osmotic water flows and refilling.

We tested the hypothesis that leaf hydraulics is correlated with the light adaptation of different plant species and specifically that the hydraulic resistance of the leaf venation (R venation ) is lower in sun- than in shade-adapted species. R venation was measured in six sun- and six shade-adapted species with a high-pressure flow meter (HPFM). The number of conduits at the proximal third of the midrib was counted and the diameter of the widest conduits together with vein density were measured. R venation was higher in shade species than in sun species and it was negatively correlated with the mean diameter of the widest conduits. Maximum leaf conductance to water vapour recorded for the different species was negatively correlated with the corresponding R venation . Sun-adapted species coping with the high water demand typical of sunny habitats appeared to have developed a highly efficient conducting system to supply living mesophyll cells with water. In contrast, species adapted to shady habitats showed higher R venation values according to their lower need for investment of carbon into producing wide conduits in the leaf.

The leaf hydraulic conductance (K leaf ) of Coffea arabica L. was measured for shoots exposed to non-lethal temperature stress or to a freeze-thaw cycle, and compared with K leaf of non-stressed samples (controls). Exposure to temperatures below 6°C for 1 h caused measurable damage to the functional integrity of cell membranes as shown by increased membrane leakiness to electrolytes. A 1 : 1 relationship was found to exist between relative electrolyte leakage and relative K leaf suggesting that membrane damage caused K leaf to increase. Low temperatures did not cause membrane disruption as shown by the comparison of chilled samples with frozen-thawed ones. In frozen leaves, membranes were extensively disrupted and both electrolyte leakiness and K leaf increased 5-fold. Low temperatures did not induce alterations of the hydraulic properties of the leaf vascular system, as revealed by measurements of K leaf after up to 500 cuttings of minor veins were made in the leaf blade of control and chilled leaves. Calculations showed that 62-75% of leaf hydraulic resistance resided in the extra-vascular water pathway. Results are discussed within the frame work of our current understanding of leaf hydraulic architecture as well as in terms of plant adaptation to extremes in temperature.