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Grapevines are being challenged by climate changes, forcing winemakers to implement irrigation systems to cope with excessive water stress. Previous studies focused on a small set of international varieties, and only few data are available for terroirs hosting cultivars with possibly different responses to drought stress. In this light, we monitored grapevine water status and grape’s physical and chemical composition, as well as concentration and structural characteristics of grape extractable polyphenols, in ten different Refosk vineyards located in the Classical Karst terroir during 2018 and 2019. Grapevines did not suffer severe stress during the two years, but their response to water shortage periods was highly heterogeneous, as pre-dawn (Ψpd) and minimum (Ψmin) leaf water potential significantly differed between vineyards, especially during the drier part of the season. Moreover, the timing of maximum water stress differed in the two years, as in 2019 longer water shortage periods and higher temperature occurred at flowering stage and before veraison, while in 2018 they were higher after veraison. These differences influenced berry’s quality, as titratable and malic acid concentration in juice, as well as total anthocyanin, total polyphenols and higher high molecular weight proanthocyanidins (HMWP) concentration in skins, were higher in 2019 than in 2018. Regarding seed proanthocyanidins, HMWP concentration, mean degree of polymerisation (mDP) and percentage of galloylation (G) in seeds were higher in 2018 than in 2019. The differences in water status measured in spatially close-related vineyards strongly support the importance of monitoring grapevines’ water status dynamics to design adequate and effective water management activities rather than relying on climate data solely. Moreover, the timing of water shortage periods also played a role in determining Refosk grape quality. Our analyses showed that the higher (but still moderate, with Ψpd and Ψmin mean values around −0.50 and −1.25 MPa, respectively) water stress between veraison and harvest occurred in 2018 might reduce Refosk grape acidity and increase concentration, polymerisation and galloylation of seed extractable proanthocyanidins.

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.

On the 29th of October 2018, a storm named “Vaia” hit North-Eastern Italy, causing the loss of 8 million m3 of standing trees and creating serious damage to the forested areas, with many economic and ecological implications. This event brought up the necessity of a standard procedure for windthrow detection and monitoring based on satellite data as an alternative to foresters’ fieldwork. The proposed methodology was applied in Carnic Alps (Friuli Venezia Giulia, NE Italy) in natural stands dominated by Picea abies and Abies alba. We used images from the Sentinel-2 mission: 1) to test vegetation indices performance in monitoring the vegetation dynamics in the short period after the storm, and 2) to create a windthrow map for the whole Friuli Venezia Giulia region. Results showed that windthrows in forests have a significant influence on visible and short-wave infrared (SWIR) spectral bands of Sentinel-2, both in the short and the long-term timeframes. NDWI8A and NDWI were the best indices for windthrow detection (R2 = 0.80 and 0.77, respectively) and NDVI, PSRI, SAVI and GNDVI had an overall good performance in spotting wind-damaged areas (R2 = 0.60–0.76). Moreover, these indices allowed to monitor post-Vaia forest die-off and showed a dynamic recovery process in cleaned sites. The NDWI8A index, employed in the vegetation index differencing (VID) change detection technique, delimited damaged areas comparable to the estimations provided by Regional Forest System (2545 ha and 3183 ha, respectively). Damaged forests detected by NDWI8A VID ranged from 500 m to 1500 m a.s.l., mainly covering steep slopes in the south and east aspects (42% and 25%, respectively). Our results suggested that the NDWI8A VID method may be a cost-effective and accurate way to produce windthrow maps, which could limit the risks associated with fieldwork and may provide a valuable tool to plan tree removal interventions in a more efficient way.

Global warming is exposing plants to increased risks of drought-driven mortality. Recent advances suggest that hydraulic failure is a key process leading to plant death, and the identification of simple and reliable proxies of species-specific risk of irreversible hydraulic damage is urgently required. We assessed the predictive power of leaf water content and shrinkage for monitoring leaf hydraulic failure in two Mediterranean native species, Salvia ceratophylloides (Sc) and S. officinalis (So). The study species showed significant differences in relative water content (RWC) thresholds inducing loss of rehydration capacity, as well as leaf hydraulic conductance (KL) impairment. Sc turned out to be more resistant to drought than So. However, Sc and So showed different leaf saturated water content values, so that different RWC values actually corresponded to similar absolute leaf water content. Our findings suggest that absolute leaf water content and leaf water potential, but not RWC, are reliable parameters for predicting the risk of leaf hydraulic impairment of two Salvia species, and their potential risk of irreversible damage under severe drought. Moreover, the lack of any KL decline until the turgor loss point in Sc, coupled to consistent leaf shrinkage, rejects the hypothesis to use leaf shrinkage as a proxy to predict KL vulnerability, at least in species with high leaf capacitance. Robust linear correlations between KL decline and electrolyte leakage measurements suggested a role of membrane damage in driving leaf hydraulic collapse.

• Increasing frequency and severity of drought events is posing risks to trees’ health, including those planted in urban settlements. Drought-induced decline of urban trees negatively affects ecosystem services of urban green spaces, and implies cost for maintenance and removal of plants. We aimed at identifying physiological traits that can explain and predict the species-specific vulnerability to climate change in urban habitats. • We assessed the relationships between long-term risk of decline of different tree species in a medium-sized town and their key indicators of drought stress tolerance i.e. turgor loss point (TLP) and vulnerability to xylem embolism (P50). • Starting from 2012, the study area experienced several summer seasons with positive anomalies of temperature and negative anomalies of precipitation. This trend was coupled with increasing percentages of urban trees showing signs of crown die-back and mortality. The species-specific risk of decline was higher for species with less negative TLP and P50 values. • The relationship between species-specific risk of climate change-induced decline of urban trees and key physiological indicators of drought tolerance confirms findings obtained in natural forests, and highlights TLP and P50 as useful indicators for species selection for tree plantation in towns, to mitigate negative impacts of climate change.

Invasive Alien Species (IAS) are a serious threat to biodiversity, severely affecting natural habitats and species assemblages. However, no consistent empirical evidence emerged on which functional traits or trait combination may foster community invasibility. Novel insights on the functional features promoting community invasibility may arise from the use of mechanistic traits, like those associated with drought resistance, which have been seldom included in trait-based studies. Here, we tested for the functional strategies of native and invasive assemblage (i.e., environmental filtering hypothesis vs. niche divergence), and we assessed how the functional space determined by native species could influence community invasibility at the edges of a resource availability gradient. Our results showed that invasive species pools need to have a certain degree of differentiation in order to persist in highly invaded communities, suggesting that functional niche divergence may foster community invasibility. In addition, resident native communities more susceptible to invasion are those which, on average, have higher resource acquisition capacity, and lower drought resistance coupled with an apparently reduced water-use efficiency. We advocate the use of a mechanistic perspective in future research to comprehensively understand invasion dynamics, providing also new insights on the factors underlying community invasibility in different ecosystems.

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.

The seedling stage is the most susceptible one during a tree′s life. Water relations may be crucial for seedlings due to their small roots, limited water buffers and the effects of drought on water transport. Despite obvious relevance, studies on seedling xylem hydraulics are scarce as respective methodical approaches are limited. Micro‐CT scans of intact Acer pseudoplatanus and Fagus sylvatica seedlings dehydrated to different water potentials (Ψ) allowed the simultaneous observation of gas‐filled versus water‐filled conduits and the calculation of percentage loss of conductivity (PLC) in stems, roots and leaves (petioles or main veins). Additionally, anatomical analyses were performed and stem PLC measured with hydraulic techniques. In A. pseudoplatanus, petioles showed a higher Ψ at 50% PLC (Ψ50 −1.13MPa) than stems (−2.51 MPa) and roots (−1.78 MPa). The main leaf veins of F. sylvatica had similar Ψ50 values (−2.26 MPa) to stems (−2.74 MPa) and roots (−2.75 MPa). In both species, no difference between root and stems was observed. Hydraulic measurements on stems closely matched the micro‐CT based PLC calculations. Micro‐CT analyses indicated a species‐specific hydraulic architecture. Vulnerability segmentation, enabling a disconnection of the hydraulic pathway upon drought, was observed in A. pseudoplatanus but not in the especially shade‐tolerant F. sylvatica. Hydraulic patterns could partly be related to xylem anatomical traits.

Plant invasive alien species (IAS) are a serious threat to biodiversity. Several studies have compared the functional features of IAS and native species to identify the functional traits, or set of traits, favouring the process of invasion. However, most of these studies analysed traits only related to carbon and nutrients, and the inclusion of traits related to water use and acquisition might be useful to describe the functions underlying plant invasion. Here we present an analysis of cost-related, hydraulic and leaf vein traits measured on a large assemblage of woody and herbaceous native and invasive species (93 species in total, 78 natives and 15 IAS), that co-occur in site types in the Mediterranean area with different water availability (i.e. xeric, mesic and hydric sites). IAS shared lower leaf construction costs and drought resistance, but potential higher efficiency in water transport (i.e. higher values of vein length per unit area/mass) than native species. Moreover, IAS and native species separated along the trade-offs drawn by the measured traits, suggesting that hydraulic and vein traits could set an important axis of variation between IAS and native species. At last, IAS tended to occupy the fast-growth region of the functional space, independently of growth form and site type. Hydraulic and vein traits provide stronger mechanistic linkages between construction costs and photosynthetic and growth rates, thus possibly playing a central role in determining the invasive potential of IAS. IAS could reduce costs associated with leaf construction and resistance to drought stress having, at the same time, high efficiency of water transport and photosynthetic rates by developing a denser venation network, translating to higher growth rates than native and more conservative species.

The choice of the best sampling strategy to capture mean values of functional traits for a species/population, while maintaining information about traits’ variability and minimizing the sampling size and effort, is an open issue in functional trait ecology. Intraspecific variability (ITV) of functional traits strongly influences sampling size and effort. However, while adequate information is available about intraspecific variability between individuals (ITVBI) and among populations (ITVPOP), relatively few studies have analyzed intraspecific variability within individuals (ITVWI). Here, we provide an analysis of ITVWI of two foliar traits, namely specific leaf area (SLA) and osmotic potential (π), in a population of Quercus ilex L. We assessed the baseline ITVWI level of variation between the two traits and provided the minimum and optimal sampling size in order to take into account ITVWI, comparing sampling optimization outputs with those previously proposed in the literature. Different factors accounted for different amount of variance of the two traits. SLA variance was mostly spread within individuals (43.4% of the total variance), while π variance was mainly spread between individuals (43.2%). Strategies that did not account for all the canopy strata produced mean values not representative of the sampled population. The minimum size to adequately capture the studied functional traits corresponded to 5 leaves taken randomly from 5 individuals, while the most accurate and feasible sampling size was 4 leaves taken randomly from 10 individuals. We demonstrate that the spatial structure of the canopy could significantly affect traits variability. Moreover, different strategies for different traits could be implemented during sampling surveys. We partially confirm sampling sizes previously proposed in the recent literature and encourage future analysis involving different traits.

Questions: 1) Are there differences in abundance-weighted functional trait values between native and alien species in coastal plant communities? 2) Which functional traits are associated with a higher level of invasion in these communities? 3) Do functional diversity patterns differ between native and alien species? 4) Is alien species occurrence linked to small-scale functional homogenization effects on the resident native species? Location: N-Adriatic coastal ecosystems (Marano and Grado lagoon, Friuli Venezia Giulia region, Italy) Methods: We sampled coastal vegetation within two habitats (foredunes and saltmarshes) along 9 belt transects in two sampling sites. Plant species richness and abundance were assessed in 128 plots along with a suite of plant functional traits. We tested for differences in CWMs between native and alien species within the two habitats, and a Linear Mixed Model (LMMs) provided insights on traits fostering the invasion success among alien species. To check for potential functional homogenization driven by alien species invasion, we explored functional diversity patterns of native and alien species (alpha and beta functional diversity) and the relationship between alpha functional diversity and alien cover. Results: Alien species had lower functional diversity than natives and were characterized by lower leaf construction costs coupled with lower drought resistance and higher water transport efficiency. The most abundant aliens were the ones minimizing carbon investment for leaf construction. In addition, we also found evidence for small-scale functional homogenization driven by alien invasion. Conclusions: Our results suggested that native species adopt a resource conservative strategy whereas alien species are characterized by a higher resource acquisition capacity (i.e. acquisitive strategy). Our data also confirmed that alien species are less functionally diverse than natives, Functional diversity of coastal plant communities potentially driving the community towards small-scale functional homogenization, resulting in a loss of species and a reduction in the functional space.

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.

Despite that congruence across taxa has been proved as an effective tool to provide insights into the processes structuring the spatial distribution of taxonomic groups and is useful for conservation purposes, only a few studies on cross-taxon congruence focused on freshwater ecosystems and on the relations among vascular plants and lichens. We hypothesized here that, since vascular plants could be good surrogates of lichens in these ecosystems, it would be possible to assess the overall biodiversity of riparian habitats using plant data only. In this frame, we explored the relationship between (a) species richness and (b) community composition of plants and lichens in a wetland area located in central Italy to (i) assess whether vascular plants are good surrogates of lichens and (ii) to test the congruence of patterns of species richness and composition among plants and lichens along an ecological gradient. The general performance of plant species richness per se, as a biodiversity surrogate of lichens, had poor results. Nonetheless, the congruence in compositional patterns between lichens and vascular plants varied across habitats and was influenced by the characteristics of the vegetation. In general, we discussed how the strength of the studied relationships could be influenced by characteristics of the data (presence/absence vs. abundance), by the spatial scale, and by the features of the habitats. Overall, our data confirm that the more diverse and structurally complex the vegetation is, the more diverse are the lichen communities it hosts.

The xylem is a complex system that includes a network of dead conduits ensuring long-distance water transport in plants. Under ongoing climate changes, xylem embolism is a major and recurrent cause of drought-induced tree mortality. Non-structural carbohydrates (NSC) play key roles in plant responses to drought and frost stress, and several studies putatively suggest their involvement in the regulation of xylem water transport. However, a clear picture on the roles of NSCs in plant hydraulics has not been drawn to date. We summarize the current knowledge on the involvement of NSCs during embolism formation and subsequent hydraulic recovery. Under drought, sugars are generally accumulated in xylem parenchyma and in xylem sap. At drought-relief, xylem functionality is putatively restored in an osmotically driven process involving wood parenchyma, xylem sap and phloem compartments. By analyzing the published data on stem hydraulics and NSC contents under drought/frost stress and subsequent stress relief, we found that embolism build-up positively correlated to stem NSC depletion, and that the magnitude of post-stress hydraulic recovery positively correlated to consumption of soluble sugars. These findings suggest a close relationship between hydraulics and carbohydrate dynamics. We call for more experiments on hydraulic and NSC dynamics in controlled and field conditions.

Invasion of natural habitats by alien trees is a threat to forest conservation. Our understanding of fundamental ecophysiological mechanisms promoting plant invasions is still limited, and hydraulic and water relation traits have been only seldom included in studies comparing native and invasive trees. We compared several leaf and wood functional and mechanistic traits in co-occurring Ailanthus altissima (Mill.) Swingle (Aa) and Fraxinus ornus L. (Fo). Aa is one of the most invasive woody species in Europe and North America, currently outcompeting several native trees including Fo. We aimed at quantifying inter-specific differences in terms of: (i) performance in resource use and acquisition; (ii) hydraulic efficiency and safety; (iii) carbon costs associated to leaf and wood construction; and (iv) plasticity of functional and mechanistic traits in response to light availability. Traits related to leaf and wood construction and drought resistance significantly differed between the two species. Fo sustained higher structural costs than Aa, but was more resistant to drought. The lower resistance to drought stress of Aa was counterbalanced by higher water transport efficiency, but possibly required mechanisms of resilience to drought-induced hydraulic damage. Larger phenotypic plasticity of Aa in response to light availability could also promote the invasive potential of the species.

Drought tolerance was greater in the whole lichen than in its isolated photobiont. Cell turgor state has an influence on the functionality of photosynthetic process in lichens. Irreversible thermodynamics is widely used to describe the water relations of vascular plants. However, poikilohydrous organisms like lichens and aeroterrestrial microalgae have seldom been studied using this approach. Water relations of lichens are generally addressed without separate analysis of the mycobiont and photobiont, and only few studies have correlated changes in photosynthetic efficiency of dehydrating lichens to accurate measurements of their water potential components. We measured water potential isotherms and chlorophyll a fluorescence in the lichen Flavoparmelia caperata harvested in different seasons, as well as in its isolated photobiont, the green alga Trebouxia gelatinosa, either exposed to water stress cycles or fully hydrated. No significant seasonal trends were observed in lichen water relations parameters. Turgor loss point and osmotic potential of the whole thallus were significantly lower than those measured in the photobiont, while differences between the water stressed photobiont and controls were not significant. Dehydration-induced drop of F v/F m was correlated with turgor loss, revealing that the photosynthetic activity of lichens partly depends on their turgor level. We provided one of the first quantitative evidences of the influence that turgor status could exert on the functionality of photosynthetic processes in lichens.

The relation between water status and expression profiles of desiccation -related genes has been studied in the desiccation tolerant (DT) aeroterrestrial green microalga Trebouxia gelatinosa, a common lichen photobiont. Algal colonies were desiccated in controlled conditions and during desiccation water content (WC) and water potential (Ψ) were measured to find the turgor loss point (Ψtlp). Quantitative real-time PCR was performed to measure the expression of ten genes related to photosynthesis, antioxidant defense, expansins, heat shock proteins (HSPs), and desiccation related proteins in algal colonies collected during desiccation when still at full turgor (WC > 6 g H2O g−1 dry weight), immediately before and after Ψtlp (−4 MPa; WC∼1 g H2O g−1 dry weight) and before and after complete desiccation (WC < 0.01 g H2O g−1 dry weight), quantifying the HSP70 protein levels by immunodetection. Our analysis showed that the expression of eight out of ten genes changed immediately before and after Ψtlp. Interestingly, the expression of five out of ten genes changed also before complete desiccation, i.e. between 0.2 and 0.01 g H2O g−1 dry weight. However, the HSP70 protein levels were not affected by changes in water status. The study provides new evidences of the link between the loss of turgor and the expression of genes related to the desiccation tolerance of T. gelatinosa, suggesting the former as a signal triggering inducible mechanisms.

Novel features related to turgor loss adjustment and higher ability to modulate leaf-construction costs could improve A. altissima fitness in its invaded range. Phenotypic plasticity has been recently proposed to play an important role in invasion process. More precisely, phenotypic plasticity of alien invasive species could be higher in their exotic rather than native habitat, probably because of the release from biotic or abiotic constraints. In this paper, several plant functional traits were analysed on Ailanthus altissima (Mill.) Swingle, a highly invasive species in Europe, aiming at providing a comparison among key functional traits measured on individuals growing in their native habitat and to test if values of functional traits and their related plasticity are higher in exotic habitats. Our analysis pinpointed that variability of functional traits in the native habitat was mainly driven by different irradiance, temperature and evaporative demand of the studied sites, in accordance to the traits trade-offs in the Leaf Economic Spectrum. Physiological traits related to drought tolerance were different between native and exotic habitats. In the native one, A. altissima relied on osmoregulation processes to adjust leaf turgor loss point (Ψtlp), as the osmotic potential at full turgor (π0) were lower in drier sites. In the exotic habitat, individuals in drier sites had similar π0 but lower wall elasticity (e), suggesting that leaves had thinner cell walls. Moreover, plasticity in specific leaf area and e were higher in the exotic habitat. The novel features related to Ψtlp adjustment and the higher plasticity in traits related to leaf-construction costs could increase A. altissima fitness in response to different environmental conditions in its invasive range.

Functional coordination between leaf and stem hydraulics has been proposed as a key trait of drought-resistant plants. A balanced water transport efficiency and safety of different plant organs might be of particular importance for plant survival in the Mediterranean climate. We monitored seasonal changes of leaf and stem water relations of Salvia officinalis L. in order to highlight strategies adopted by this species to survive in harsh environmental conditions. During summer drought, the water potential dropped below the turgor loss point thus reducing water loss by transpiration, whereas the photosynthetic efficiency remained relatively high. Leaves lost their water transport efficiency earlier than stems, although in both plant organs P50 (water potential inducing 50% loss of hydraulic conductivity) indicated surprisingly high vulnerability when compared with other drought-tolerant species. The fast recovery of leaf turgor upon restoration of soil water availability suggests that the reduction of leaf hydraulic conductance is not only a consequence of vein embolism, but cell shrinkage and consequent increase of resistance in the extra-xylem pathway may play an important role. We conclude that the drought tolerance of S. officinalis arises at least partly as a consequence of vulnerability segmentation.