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Understanding which structural and functional traits are linked to species’ vulnerability to embolism formation (P50) may provide fundamental knowledge on plant strategies to maintain an efficient water transport. We measured P50, wood density (WD), mean conduit area, conduit density, percentage areas occupied by vessels, parenchyma cells (PATOT) and fibers (FA) on branches of angiosperm and gymnosperm species. Moreover, we compiled a dataset of published hydraulic and anatomical data to be compared with our results. Species more vulnerable to embolism had lower WD. In angiosperms, the variability in WD was better explained by PATOT and FA, which were highly correlated. Angiosperms with a higher P50 (less negative) had a higher amount of PATOT and total amount of nonstructural carbohydrates. Instead, in gymnosperms, P50 vs PATOT was not significant. The correlation between PATOT and P50 might have a biological meaning and also suggests that the causality of the commonly observed relationship of WD vs P50 is indirect and dependent on the parenchyma fraction. Our study suggests that angiosperms have a potential active embolism reversal capacity in which parenchyma has an important role, while in gymnosperms this might not be the case.

Stem photosynthesis has been suggested to play relevant roles to cope with different biotic and abiotic stress factors, including drought. In the present study, we performed measurements of stem hydraulic conductance and non-structural carbohydrate content in the evergreen Laurus nobilis L. and the deciduous Populusalba L., subjected to inhibition of stem photosynthesis and successive exposure to a drought-recovery cycle in order to check if stem photosynthesis may be involved in allowing hydraulic recovery after drought stress relief. Stem shading affected the growth of L. nobilis but not of P. alba saplings. By contrast, inhibition of stem photosynthesis was coupled to inhibition of hydraulic recovery following embolism build-up under drought in P. alba but not in L. nobilis. The two study species showed a different content and behavior of nonstructural carbohydrates (NSCs). The differences in NSCs’ trend and embolism reversal ability led to a significant relationship between starch content and the corresponding hydraulic conductance values in L. nobilis but not in P. alba. Our findings suggest that stem photosynthesis plays a key role in the maintenance of hydraulic functioning during drought especially in the deciduous species. This, in turn, may increase their vulnerability under current global climate change scenarios.

Accurate predictions of species distribution under current and future climate conditions require modeling efforts based on clear mechanistic relationships between climate variables and plant physiological functions. Vulnerability of leaves to xylem embolism is a key mechanistic trait that might be included in these modeling efforts. Here, we propose a simple set-up to measure leaf vulnerability to embolism on the basis of the optical method using a smartphone, a light source, and a notebook. Our data show that this proposed set-up can adequately quantify the vulnerability to xylem embolism of leaf major veins in Populus nigra and Ostrya carpinifolia, producing values consistent with those obtained in temperate tree species with other methods, allowing virtually any laboratory to quantify species-specific drought tolerance on the basis of a sound mechanistic trait.

Embolism repair ability has been documented in numerous species. Although the actual mechanism driving this phenomenon is still debated, experimental findings suggest that non-structural carbohydrates (NSC) stored in wood parenchyma would provide the osmotic forces to drive the refilling of embolized conduits. We selected 12 broadleaved species differing in vulnerability to xylem embolism (P-50) and amount of wood parenchyma in order to check direct evidence about the possible link(s) between parenchyma cells abundance, NSC availability and species-specific capacity to reverse xylem embolism. Branches were dehydrated until similar to 50% loss of hydraulic conductivity was recorded (PLC similar to 50%). Hydraulic recovery (Delta PLC) and NSC content was, then, assessed after 111 of rehydration. Species showed a different ability to recover their hydraulic conductivity from PLC similar to 50%. Removing the bark in the species showing hydraulic recovery inhibited the embolism reversal. Strong correlations between the Delta PLC and: a) the amount of parenchyma cells (mainly driven by the pith area), b) the consumption of soluble NSC have been recorded. Our results support the hypothesis that refilling of embolized vessels is mediated by the mobilization of soluble NSC and it is mainly recorded in species with a higher percentage of parenchyma cells that may be important in the hydraulic recovery mechanism as a source of carbohydrates and/or as a source of water.

The maintenance of hydraulic function during and after a drought event is crucial for tree survival, but the importance of non-structural carbohydrates (NSCs) in the recovery phase is still debated. We tested whether higher NSC availability facilitates post-drought hydraulic recovery, by applying a short-term drought (Sdr) and a long-term drought combined with shading (Ldr+sh) in Fraxinus ornus and Ostrya carpinifolia. Plants were then re-irrigated and recovery was checked 24 h later, by measuring water potential, stem percentage loss of hydraulic conductance (PLC) and NSC content. The relative magnitude of hydraulic and carbon constraints was also assessed in desiccated plants. During drought, PLC increased only in F. ornus, while it was maintained almost constant in O. carpinifolia due to tighter stomatal control of xylem pressure (i.e. more isohydric). In F. ornus, only Sdr plants maintained high NSC contents at the end of drought and, when re-irrigated, recovered PLC to control values. Whereas hydraulic failure was ubiquitous, only F. ornus depleted NSC reserves at mortality. Our results suggest that preserving higher NSC content at the end of a drought can be important for the hydraulic resilience of trees.

International audience; Three woody species typically encroaching the Karstic region of Trieste (Northeastern Italy) i.e. Cotinus coggygria L., Prunus mahaleb L. and Fraxinus ornus L., have been measured for diurnal and seasonal time courses of leaf conductance to water vapour, transpiration and water potential ($\Psi _{\rm L}$), as well as whole-plant, shoot and root hydraulic resistance (R$_{{\rm plant}}$, R$_{{\rm shoot}}$ and R$_{{\rm root}}$, respectively) in view of investigating the impact of plant hydraulics on water stress resistance. F. ornus suffered the highest $\Psi _{\rm L}$ drop in July and September due to the high Rplant measured in this species. Positive significant relations were found of R$_{{\rm plant}}$ and $\Psi _{\rm L}$ to R$_{{\rm root}}$ in all three species studied, thus suggesting that root hydraulics is a major determinant of plant hydraulics and water stress resistance. The increasing R$_{{\rm root}}$ (and R$_{{\rm plant}}$) from C. coggygria to P. mahaleb to F. ornus provided a possible explanation for the typical temporal sequence of expansion of the three species into degraded areas.; Relations hydriques et caractéristiques hydrauliques de trois espèces ligneuses présentes dans le même habitat. Trois espèces ligneuses, typiquement en cours d'extension dans la région Karstique de Trieste (Italie du nord) i.e. Cotinus coggygria L., Prunus mahaleb L. and Fraxinus ornus L., ont été suivies aux échelles journalières et saisonnières en termes de conductance foliaire pour la vapeur d'eau, de transpiration et de potentiel hydrique ($\Psi _{\rm L}$), ainsi que de résistance hydraulique totale, tige et racinaire (R$_{{\rm plant}}$, R$_{{\rm shoot}}$ and R$_{{\rm root}}$, respectivement) dans l'objectif d'analyser l'impact des paramètres hydrauliques des plantes sur leur résistance au stress hydrique. F. ornus a montré la plus grande chute de $\Psi _{\rm L}$ en Juillet et Septembre en raison de la grande valeur de R$_{{\rm plant}}$ mesurée chez cette espèce. Des relations positives et significatives ont été trouvées entre R$_{{\rm plant}}$, $\Psi _{\rm L}$ et R$_{{\rm root}}$ pour les trois espèces étudiées, ce qui suggère que les propriétés hydrauliques des racines sont un déterminant majeur des propriétés hydrauliques de la plante et de leur résistance au stress hydrique. Les valeurs croissantes de R$_{{\rm root}}$ (et R$_{{\rm plant}}$) de C. coggygria à P. mahaleb et F. ornus fournissent une explication possible pour la séquence temporelle d'expansion de ces trois espèces dans les zones dégradées.

• The hydraulic conductance of angiosperm xylem has been suggested to vary with changes in sap solute concentrations because of intervessel pit properties. • The magnitude of the ‘ionic effect’ was linked with vessel and pit dimensions in 20 angiosperm species covering 13 families including six Lauraceae species. • A positive correlation was found between ionic effect and vessel grouping parameters, especially the portion of vessel walls in contact with neighbouring vessels. Species with intervessel contact fraction (FC) values < 0.1 showed an ionic effect between 2% and 17%, while species with FC values > 0.1 exhibited a response between 10% and 32%. The ionic effect increased linearly with the mean fraction of the total vessel wall area occupied by intervessel pits as well as with the intervessel contact length. However, no significant correlation occurred between the ionic effect and total intervessel pit membrane area per vessel, vessel diameter, vessel length, vessel wall area, and intervessel pit membrane thickness. • Quantitative vessel and pit characters are suggested to contribute to interspecific variation of the ionic effect, whereas chemical properties of intervessel pit membranes are likely to play an additional role.

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 (Px) 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 Px, with HSC-cells between 65 and 75% of the total at Px ≥ –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 Px 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 Px 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.