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Mistletoes are xylem-tapping hemiparasites that rely on their hosts for water and nutrient uptake. Thus, they impair tree performance in the face of environmental stress via altering the carbon and water relations and nutritional status of trees. To improve our understanding of physiological responses to mistletoe and ongoing climate change, we investigated radial growth, stable carbon and oxygen isotopic signals, and elemental composition of tree rings in silver fir (Abies alba Mill.) and Scots pine (Pinus sylvestris L.) forests infested with Viscum album L. We compared temporal series (1990-2020) of basal area increment (BAI), intrinsic water-use efficiency (iWUE), oxygen isotope composition (δ18O), nutrient concentrations and stoichiometric ratios between non-infested (NI) and severely infested (SI) fir and pine trees from populations located close to the xeric distribution limit of the species in north-eastern Spain. The SI trees showed historically higher growth, but the BAI trend was negative for more than three decades before 2020 and their growth rates became significantly lower than those of NI trees by the mid-2010s. Mistletoe infestation was related to an enhanced sensitivity of radial growth to vapour pressure deficit (atmospheric drought). The SI trees showed less pronounced iWUE increases (fir) and lower iWUE values (pine) than NI trees. The lower tree-ring δ18O values of SI trees may be the result of several superimposed effects operating simultaneously, including leaf-level evaporative enrichment, source water isotopic signals, and anatomical and phenological differences. We observed a deterioration of potassium (K) nutrition in tree-ring wood of both species in SI trees, along with accumulation of manganese (Mn). We suggest that such nutritional patterns are driven by the indirect effect of mistletoe-induced drought stress, particularly in pine. The combined analyses of different physiological indicators imprinted on tree rings provided evidence of the progressive onset of carbon, water and nutrient imbalances in mistletoe-infested conifers inhabiting seasonally dry regions., (© The Author(s) 2024. Published by Oxford University Press.)

Coastal dunes are characterised by strong gradients of abiotic stress, typically increasing in severity from inland areas towards the shoreline. Thus, dune gradients represent unique opportunities to study intraspecific responses to environmental changes and to investigate which factors drive community change. This study aims to examine functional trait variation in two coexisting species in response to environmental changes along a dune gradient in NW Spain. Trait convergence was also investigated and compared between both ends of the gradient. We measured functional leaf traits related to plant efficiency in the use of light, water and nutrients, also possible stressors (salt content and pH) and availability of limiting resources (water and nutrients) in the soil. Most soil variables showed changes following a non-directional gradient. Differences in soil variables were site specific and depended on growth of the study species. Structural and functional traits depended on species and/or plant position on the gradient, except for effective quantum yield of PSII and leaf δ 15 N. The pattern of variation was mostly directional for reflectance indices related to leaf physiology. Multivariate analyses showed significant interspecific differences in the set of traits they exhibited along positions in the gradient. Species also differed in the combination of traits selected under given environmental conditions. Coexisting species display a specific set of traits that reflects different strategies to environmental stress. Our study highlights the overly simplistic nature of some previous studies that assume dune gradients are monotonically directional, without considering that these gradients may be differentially modified by species activity., (© 2024 The Author(s). Plant Biology published by John Wiley & Sons Ltd on behalf of German Society for Plant Sciences, Royal Botanical Society of the Netherlands.)

Access to extractive resources relies as much on technology and prices as it does on the power to designate ecological and economic meanings to water and other environmental goods. This paper examines the ways in which the mining industry uses scientific models to create meanings for water that in turn legitimizes their access to and control over it. To do so, this paper explores the relevance of combining biophysical analyses-in this case water metabolism-with an examination of those power relations and social constructions that coexist with and affect the flows of water. Based on empirical research, this paper analyzes the evolution of water management in the process of copper production at the Cobre las Cruces mine in southern Spain to identify present contradictions in the strategies adopted by the mining company to avoid water degradation. These contradictions are revealed by examining how water has been framed as a resource not susceptible to being used for purposes other than mining processes. We argue that those framing this environmental explanation-the regional government and the mining industry-are promoting net subtractions of water from an aquifer against current regulations. [ABSTRACT FROM AUTHOR]

Three freshwater microalgae (Spirogyra sp., Cosmarium sp., and Cosmarium blytii) collected from several locations in Gran Canaria have been studied to explore their potential as a novel source of bioactive compounds for biotechnological applications. Soluble carbohydrates were quantified after extraction with 3 M HCl at 100 °C, ranging from 35.8 to 43.3 %, and with water at room temperature, ranging from 19 to 22.8 %. Amino acids glutamic acid, proline and aspartic acid were quantified by RP-HPLC. Glutamic acid was the most abundant, ranging from 12.2 to 3.63 mg g -1 of dry biomass. Cosmarium blytii was the richest sample in amino acids (24.02 mg g -1 of dry weight). In addition, Cosmarium blytii and Spyrogira sp. exhibited higher radical scavenging activity (RSA) against 1,1-diphenyl-2-picrylhydrazyl (DPPH) than that of the synthetic antioxidant butylhydroxytoluene (BHT), commonly used as food additive. These results show a great potential of these microalgae for exploitation in the food, feed and pharmaceutical industries., (© 2022 The Authors. Chemistry & Biodiversity published by Wiley-VHCA AG, Zurich, Switzerland.)

Human societies and ecosystems use water in different ways and at different scales, which complicates the study of water-use sustainability in socioecological systems. We present a multiscale integrated assessment of societal and ecosystem metabolism, an innovative approach to the quantitative analysis of water use that addresses the problem of multiple scales. It builds on the concept of metabolic pattern and the flow -- fund model of Georgescu-Roegen. We show how to define water resources and water use (expressed in hourly rates) for socioeconomic systems in relation to the identities of relevant fund elements (relevant categories of human activity or land use) over a time span of 1 year. Similarly, we define the limits on the human appropriation of water (aggregate withdrawal or damping per year) on the basis of the structural and functional stability of ecological funds (defined over a much longer time scale) and the related land-use pattern. [ABSTRACT FROM AUTHOR]

Despite the importance of understanding plant growth, the mechanisms underlying how plant and fruit growth declines during drought remain poorly understood. Specifically, it remains unresolved whether carbon or water factors are responsible for limiting growth as drought progresses. We examine questions regarding the relative importance of water and carbon to fruit growth depending on the water deficit level and the fruit growth stage by measuring fruit diameter, leaf photosynthesis, and a proxy of cell turgor in olive (Olea europaea). Flow cytometry was also applied to determine the fruit cell division stage. We found that photosynthesis and turgor were related to fruit growth; specifically, the relative importance of photosynthesis was higher during periods of more intense cell division, while turgor had higher relative importance in periods where cell division comes close to ceasing and fruit growth is dependent mainly on cell expansion. This pattern was found regardless of the water deficit level, although turgor and growth ceased at more similar values of leaf water potential than photosynthesis. Cell division occurred even when fruit growth seemed to stop under water deficit conditions, which likely helped fruits to grow disproportionately when trees were hydrated again, compensating for periods with low turgor. As a result, the final fruit size was not severely penalized. We conclude that carbon and water processes are able to explain fruit growth, with importance placed on the combination of cell division and expansion. However, the major limitation to growth is turgor, which adds evidence to the sink limitation hypothesis., (© 2021 The Authors. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Background: Water scarcity is currently affecting many areas of the world, reaching worrying levels in drought areas such as southern Spain. To cope with this issue, researchers in the agricultural sector have implemented deficit irrigation strategies intended to reduce water consumption by increasing fruit quality. Almond is among the most popular tree nuts worldwide and also the most nut cultivated in Spain. Almond consumption, together with other nuts, has been widely associated with improvements in cardiovascular health, metabolic syndrome and diabetes owing to their bioactive compounds such as polyphenols. Water deficit strategies generate hydroSOStainable almonds, raised under water stress conditions, with high content of bioactive compounds. The aim of this work was to study the relationship between water stress, color and polyphenols in hydroSOStainable almonds. For this, instrumental color, total phenolic content and phenolic compounds were measured and correlated using Pearson's correlation., Results: The results showed a strong relationship between water stress, color and polyphenols of almonds, showing that increasing water stress in plants up to ~100 MPa × day values of stress integral increase the polyphenols in almonds, leading to a reddish color., Conclusion: Finally, this research demonstrated that implementing water-saving strategies help to improve the phenolic content and color of hydroSOStainable almonds and also that isorhamnetin-3-O-rutinoside, isorhamnetin-3-O-glucoside and kaempferol-3-O-glucoside could be important markers of hydroSOStainable almonds (cv. Vairo). Besides, hydroSOStainable almonds could be an important source of phenols, providing 25% of the estimated total polyphenolic daily intake. © 2020 Society of Chemical Industry., (© 2020 Society of Chemical Industry.)

Background: Water deficit and iron deficiency (iron chlorosis) are common environmental stresses that affect grapevine production in the Mediterranean area. Studies on the impact of both stresses, when they act simultaneously, are rare. The main objective of this investigation was to evaluate the combined effects of the incidence of iron chlorosis and the vine water status on quality of Tempranillo wine. For this, 20 non-irrigated vineyard subzones (10 m × 10 m each), from non-affected to moderately affected by iron chlorosis, were monitored in the Ribera del Duero area (north-central Spain) during two consecutive seasons., Results: Factorial analyses of variance were performed to study the effects of predawn leaf water potential and foliar chlorophyll content, both measured at veraison, on chemical and sensory characteristics of wine. With an impact much greater than water status, the incidence of iron stress decreased pH of the wine and enhanced sensory attributes as tonality, layer intensity, flavour intensity, and persistence in the mouth. There were increases in red colour, astringency, and persistence of the wine associated with chlorosis, although they might be restricted in water-deficit conditions., Conclusion: The results have demonstrated that mild to moderate iron stress can have positive effects on chemical and sensory attributes of Tempranillo wine. Measurements of foliar chlorophyll content at veraison could be very useful to map quality potential in rainfed vineyards affected by iron deficiency. © 2020 Society of Chemical Industry., (© 2020 Society of Chemical Industry.)

Introduction: Water is an essential nutrient for life and the most abundant component in the human body. However, its dietary recommendations or clinical management guidelines do not receive as much attention as they deserve. In addition, there are some obstacles to establishing optimal values, both for the amount of water the body must contain and for water ingestion. Water intake and elimination depend on unsteady factors that are difficult to measure and, at the same time, compensated by the body's ability to regulate homeostasis. Since scientific evidence is lacking for establishing recommendations, "adequate intakes" (to maintain an adequate hydration state) have been estimated using data on water intake from groups of healthy people. The European Food Safety Authority (EFSA) also considers desirable the use of urine osmolarity to estimate the adequacy of water intake in adults. Clinical studies have generally shown the benefits of adequate hydration and the damage caused by water imbalance, whether quantitative (dehydration and overhydration) or qualitative (extracellular and intracellular water). Unfortunately, these studies are few and often have poor cross-sectional, case-control, or prospective designs, and use small samples or indirect methods to assess hydration status. This article presents up-to-date information on subjects such as: 1) compliance with water consumption recommendations and suggestions for improvement; 2) techniques available to measure hydration status and their clinical applications; 3) effects of hydration/dehydration on physical or cognitive activities and chronic diseases; and 4) existing Spanish regulations on the quality and salubrity of water.

The extent to which water availability can be used to predict the enlargement and final dimensions of xylem conduits remains an open issue. We reconstructed the time course of tracheid enlargement in Pinus sylvestris trees in central Spain by repeated measurements of tracheid diameter on microcores sampled weekly during a 2 yr period. We analyzed the role of water availability in these dynamics empirically through time-series correlation analysis and mechanistically by building a model that simulates daily tracheid enlargement rate and duration based on Lockhart's equation and water potential as the sole input. Tracheid enlargement followed a sigmoid-like time course, which varied intra- and interannually. Our empirical analysis showed that final tracheid diameter was strongly related to water availability during tracheid enlargement. The mechanistic model was calibrated and successfully validated (R 2  = 0.92) against the observed tracheid enlargement time course. The model was also able to reproduce the seasonal variations of tracheid enlargement rate, duration and final diameter (R 2  = 0.84-0.99). Our results support the hypothesis that tracheid enlargement and final dimensions can be modeled based on the direct effect of water potential on turgor-driven cell expansion. We argue that such a mechanism is consistent with other reported patterns of tracheid dimension variation., (© 2019 The Authors New Phytologist © 2019 New Phytologist Trust.)

Reducing water consumption on agriculture is a worldwide goal toward sustainability. In this scenario, two experiments of regulated deficit irrigation (RDI) were carried out on olive trees, cultivar Manzanilla. With regard to experiment A, three RDI techniques were applied during the olive pit hardening period (stage II), while in experiment B, two RDI treatments were applied during the rehydration phase (stage III). Table olives under RDI are so-called hydroSOStainable. The effect of water deficit and Spanish-style processing was studied on the polyphenol profile, antioxidant capacity, and total polyphenol content (TPC) of both raw olives (RO) and table olives (TO). The TPC decreased after processing of TO. It could be due to osmotic mechanisms. However, many individual polyphenols, such as oleuropein (main polyphenol) or oleoside diglucoside, increased their concentrations in hydroSOStainable TO. Additionally, the TPC content was correlated to the phenological stage of the fruit when the stress is applied. A moderate stress during pit hardening and an intense stress during the rehydration phase were the treatments that best improved the polyphenol profile.

Climatic dryness imposes limitations on vascular plant growth by reducing stomatal conductance, thereby decreasing CO 2 uptake and transpiration. Given that transpiration-driven water flow is required for nutrient uptake, climatic stress-induced nutrient deficit could be a key mechanism for decreased plant performance under prolonged drought. We propose the existence of an "isohydric trap," a dryness-induced detrimental feedback leading to nutrient deficit and stoichiometry imbalance in strict isohydric species. We tested this framework in a common garden experiment with 840 individuals of four ecologically contrasting European pines (Pinus halepensis, P. nigra, P. sylvestris, and P. uncinata) at a site with high temperature and low soil water availability. We measured growth, survival, photochemical efficiency, stem water potentials, leaf isotopic composition (δ 13 C, δ 18 O), and nutrient concentrations (C, N, P, K, Zn, Cu). After 2 years, the Mediterranean species Pinus halepensis showed lower δ 18 O and higher δ 13 C values than the other species, indicating higher time-integrated transpiration and water-use efficiency (WUE), along with lower predawn and midday water potentials, higher photochemical efficiency, higher leaf P, and K concentrations, more balanced N:P and N:K ratios, and much greater dry-biomass (up to 63-fold) and survival (100%). Conversely, the more mesic mountain pine species showed higher leaf δ 18 O and lower δ 13 C, indicating lower transpiration and WUE, higher water potentials, severe P and K deficiencies and N:P and N:K imbalances, and poorer photochemical efficiency, growth, and survival. These results support our hypothesis that vascular plant species with tight stomatal regulation of transpiration can become trapped in a feedback cycle of nutrient deficit and imbalance that exacerbates the detrimental impacts of climatic dryness on performance. This overlooked feedback mechanism may hamper the ability of isohydric species to respond to ongoing global change, by aggravating the interactive impacts of stoichiometric imbalance and water stress caused by anthropogenic N deposition and hotter droughts, respectively., (© 2018 John Wiley & Sons Ltd.)

It has been known for a long time that drought intensity is a critical variable in determining water stress of Mediterranean tree species. However, not as much attention has been paid to other drought characteristics, for example the timing of the dry periods. We investigated the impact of the timing and intensity of extreme droughts on growing season length, growth and water-use efficiency of three tree species, Pinus nigra ssp. Salzmannii J.F. Arnold, Quercus ilex ssp. ballota (Desf.) Samp. and Quercus faginea Lam. coexisting in a continental Mediterranean ecosystem. Over the study period (2009-13), intense droughts were observed at annual and seasonal scales, particularly during 2011 and 2012. In 2012, an atypically dry winter and spring was followed by an intense summer drought. Quercus faginea growth was affected more by drought timing than by drought intensity, probably because of its winter-deciduous leaf habit. Pinus nigra showed a lower decrease in secondary growth than observed in the two Quercus species in extremely dry years. Resilience to extreme droughts was different among species, with Q. faginea showing poorer recovery of growth after very dry years. The highest intra- and inter-annual plasticity in water-use efficiency was observed in P. nigra, which maintained a more water-saving strategy. Our results revealed that the timing of extreme drought events can affect tree function to a larger extent than drought intensity, especially in deciduous species. Legacy effects of drought over months and years significantly strengthened the impact of drought timing and intensity on tree function.

In Mediterranean mountains, Pinus sylvestris L. is expected to be displaced under a warming climate by more drought-tolerant species such as the sub-Mediterranean Quercus pyrenaica Willd. Understanding how environmental factors drive tree physiology and phenology is, therefore, essential to assess the effect of changing climatic conditions on the performance of these species and, ultimately, their distribution. We compared the cambial and leaf phenology and leaf gas exchange of Q. pyrenaica and P. sylvestris at their altitudinal boundary in Central Spain and assessed the environmental variables involved. Results indicate that P. sylvestris cambial phenology was more sensitive to weather conditions (temperature at the onset and water deficit at the end of the growing season) than Q. pyrenaica. On the other hand, Q. pyrenaica cambial and leaf phenology were synchronized and driven by photoperiod and temperatures. Pinus sylvestris showed lower photosynthetic nitrogen-use efficiency and higher intrinsic water-use efficiency than Q. pyrenaica as a result of a tighter stomatal control in response to summer dry conditions, despite its less negative midday leaf water potentials. These phenological and leaf gas exchange responses evidence a stronger sensitivity to drought of P. sylvestris than that of Q. pyrenaica, which may therefore hold a competitive advantage over P. sylvestris under the predicted increase in recurrence and intensity of drought events. On the other hand, both species could benefit from warmer springs through an advanced phenology, although this effect could be limited in Q. pyrenaica if it maintains a photoperiod control over the onset of xylogenesis.

Nowadays, irrigation with low quality water is becoming an alternative to satisfy the needs of crops. However, some plant species have to deal with high salinity of reclaimed water, by adapting their physiological behaviour during both saline and recovery periods and developing morphological changes in their leaves. The application of arbuscular mycorrhizal fungi (AMF) could also be a suitable option to mitigate the negative effects of this kind of water, although the effectiveness of plant-AMF association is influenced by many factors. In this work, during forty weeks, the combined effect of Glomus iranicum var. tenuihypharum and two types of water: control, C, EC<0.9 dS m(-1) and reclaimed water, RW (with EC: 4 dS m(-1) during a first saline period and EC: 6 dS m(-1) during a second saline period) was evaluated for laurustinus plants (Viburnum tinus L.) transplanted in soil. This was followed by a recovery period of eight weeks, when all the plants were irrigated in the control irrigation conditions. Seasonal and daily changes in stem water potential (Ψstem), stomatal conductance (gs), photosynthesis (Pn) and leaf internal CO2 concentration (Ci) of laurustinus plants were evaluated. Leaf structure alterations, nutrient imbalance, height and leaf hydraulic conductivity (Kleaf) were also determined. Due to the high difficulty of absorbing water from the soil, RW plants showed a high volumetric water content (θv) in soil. The stem water potential and the stomatal conductance (gs) values were reduced in RW plants throughout the second saline period. These decreases were also found during the day. Leaf Ca(2+)/Na(+) and K(+)/Na(+) ratios diminished in RW plants respect to the C plants due to the Na(+) accumulation, although height and chlorophyll content values did not show statistical differences. Leaves from RW plants showed a significantly thicker mesophyll than Control leaves as a consequence of high EC. The area of palisade parenchyma (PP) increased while the area of spongy parenchyma (SP) decreased in RW leaves with respect to the C leaves. These structural changes could be considered as a strategy to maximize photosynthesis potential in saline conditions. Mycorrhizal inoculation improved the water status of both C and RW plants by increasing their Ψstem and gs values. As regards leaf structure, AMF showed an opposite effect to salinity for PP and SP. At the end of the recovery period, hardly any statistical differences of physiological parameters were found between treatments, although a tendency to improve them was observed in inoculated plants. In any case, the leaf structural changes and the great reduction in Kleaf observed at Ψleaf below -1.5 MPa would constitute an important mechanism for laurustinus plants to reduce the water loses produced by salinity., (Copyright © 2015 Elsevier GmbH. All rights reserved.)

Background: In table grapes, berry firmness influences consumer acceptance so it is important to avoid berry shattering and dehydration during their post-harvest life. Since studies of irrigation effects on table grape quality are comparatively rare, sensory evaluation aimed to identify high-quality berries obtained under different deficit irrigation treatments. A 3-year study examined the effects of deficit irrigation strategies on some physical quality attributes at harvest, after 28 days of cold storage at 0 °C and after an additional shelf-life period of 3 days at 15 °C. Control vines were irrigated to ensure non-limiting water conditions (110% of crop evapo-transpiration), while both regulated deficit irrigation treatment (RDI) and partial root-zone drying (PRD) treatments applied 35% less water post-veraison. The null irrigation treatment (NI) only received natural precipitation (72% less water than control vines)., Results: Total yield and physical quality at harvest were not significantly affected by RDI or PRD. Only severe deficit (NI) decreased berry size, and this treatment had the most dehydrated berries and the worst sensory scores post-harvest. After cold storage, increased berry shattering of the PRD treatment was correlated with lower leaf xylem abscisic acid (ABA) concentration at the time of harvest. Overall quality, especially stem browning, determined the shelf-life, and longer storage duration tended to diminish treatment differences., Conclusions: Only NI clusters showed lower quality than their irrigated counterparts. Neither RDI nor PRD had any noticeable effect on berry quality at the end of cold storage and shelf-life, with the slight differences detected between these treatments related to stem browning and dehydration. Sensory results were similar in RDI and PRD, which provided grapes that were more acceptable to consumers than the control. Thus, it is possible to decrease irrigation of table grapes without adversely affecting the physical quality of the berries., (© 2014 Society of Chemical Industry.)

Wild fungi play a critical role in forest ecosystems, and its recollection is a relevant economic activity. Understanding fungal response to climate is necessary in order to predict future fungal production in Mediterranean forests under climate change scenarios. We used a 15-year data set to model the relationship between climate and epigeous fungal abundance and productivity, for mycorrhizal and saprotrophic guilds in a Mediterranean pine forest. The obtained models were used to predict fungal productivity for the 2021-2080 period by means of regional climate change models. Simple models based on early spring temperature and summer-autumn rainfall could provide accurate estimates for fungal abundance and productivity. Models including rainfall and climatic water balance showed similar results and explanatory power for the analyzed 15-year period. However, their predictions for the 2021-2080 period diverged. Rainfall-based models predicted a maintenance of fungal yield, whereas water balance-based models predicted a steady decrease of fungal productivity under a global warming scenario. Under Mediterranean conditions fungi responded to weather conditions in two distinct periods: early spring and late summer-autumn, suggesting a bimodal pattern of growth. Saprotrophic and mycorrhizal fungi showed differences in the climatic control. Increased atmospheric evaporative demand due to global warming might lead to a drop in fungal yields during the 21st century., (© 2015 John Wiley & Sons Ltd.)

Lichens are adapted to desiccation/rehydration and accumulate heavy metals, which induce ROS especially from the photobiont photosynthetic pigments. Although their mechanisms of abiotic stress tolerance are still to be unravelled, they seem related to symbionts' reciprocal upregulation of antioxidant systems. With the aim to study the effect of Pb on oxidative status during rehydration, the kinetics of intracellular ROS, lipid peroxidation and chlorophyll autofluorescence of whole Ramalina farinacea thalli and its isolated microalgae (Trebouxia TR1 and T. TR9) was recorded. A genetic characterization of the microalgae present in the thalli used was also carried out in order to assess possible correlations among the relative abundance of each phycobiont, their individual physiological responses and that of the entire thallus. Unexpectedly, Pb decreased ROS and lipid peroxidation in thalli and its phycobionts, associated with a lower chlorophyll autofluorescence. Each phycobiont showed a particular pattern, but the oxidative response of the thallus paralleled the TR1's, agreeing with the genetic identification of this strain as the predominant phycobiont. We conclude that: (1) the lichen oxidative behaviour seems to be modulated by the predominant phycobiont and (2) Pb evokes in R. farinacea and its phycobionts strong mechanisms to neutralize its own oxidant effects along with those of rehydration.

Vegetation in water-limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species-specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long-term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long-term experimental drought shifted water uptake toward deeper (10-35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought-affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions., (© 2014 John Wiley & Sons Ltd.)

In order to understand the impact of drought and intrinsic water-use efficiency (iWUE) on tree growth, we evaluated the relative importance of direct and indirect effects of water availability on secondary growth and xylem anatomy of Juniperus thurifera, a Mediterranean anisohydric conifer. Dendrochronological techniques, quantitative xylem anatomy, and (13)C/(12)C isotopic ratio were combined to develop standardized chronologies for iWUE, BAI (basal area increment), and anatomical variables on a 40-year-long annually resolved series for 20 trees. We tested the relationship between iWUE and secondary growth at short-term (annual) and long-term (decadal) temporal scales to evaluate whether gains in iWUE may lead to increases in secondary growth. We obtained a positive long-term correlation between iWUE and BAI, simultaneously with a negative short-term correlation between them. Furthermore, BAI and iWUE were correlated with anatomical traits related to carbon sink or storage (tracheid wall thickness and ray parenchyma amount), but no significant correlation with conductive traits (tracheid lumen) was found. Water availability during the growing season significantly modulated tree growth at the xylem level, where growth rates and wood anatomical traits were affected by June precipitation. Our results are consistent with a drought-induced limitation of tree growth response to rising CO2, despite the trend of rising iWUE being maintained. We also remark the usefulness of exploring this relationship at different temporal scales to fully understand the actual links between iWUE and secondary growth dynamics.

Winter-drought induced forest diebacks in the low-latitude margins of species' distribution ranges can provide new insights into the mechanisms (carbon starvation, hydraulic failure) underlying contrasting tree reactions. We analysed a winter-drought induced dieback at the Scots pine's southern edge through a dual-isotope approach (Δ(13) C and δ(18) O in tree-ring cellulose). We hypothesized that a differential long-term performance, mediated by the interaction between CO(2) and climate, determined the fates of individuals during dieback. Declining trees showed a stronger coupling between climate, growth and intrinsic water-use efficiency (WUEi) than non-declining individuals that was noticeable for 25 years prior to dieback. The rising stomatal control of water losses with time in declining trees, indicated by negative Δ(13) C-δ(18) O relationships, was likely associated with their native aptitude to grow more and take up more water (suggested by larger tracheid lumen widths) than non-declining trees and, therefore, to exhibit a greater cavitation risk. Freeze-thaw episodes occurring in winter 2001 unveiled such physiological differences by triggering dieback in those trees more vulnerable to hydraulic failure. Thus, WUEi tightly modulated growth responses to long-term warming in declining trees, indicating that co-occurring individuals were differentially predisposed to winter-drought mortality. These different performances were unconnected to the depletion of stored carbohydrates., (© 2013 John Wiley & Sons Ltd.)

No information exists on the proanthocyanidin content of pear-jujube (Ziziphus jujuba Mill) fruit, their polymeric types and sizes, and their self-aggregation, or on the effect of different water deficit levels during the fruit maturation period on these compounds. Two trimers, two tetramers, and six B type procyanidin pentamers were identified and quantified for the first time. Water deficit increased the content of procyanidins of low molecular mass, improving their potential bioavailability and possible physiological effects on human health. The tendency of procyanidins to self-aggregate was similar in the edible portion and pit, and was not affected by water deficit. The procyanidin content of fruit from well watered trees increased during domestic cold storage, whereas the fruits from trees suffering severe water stress lost some of their procyanidin content.

We studied the intraspecific variability of maritime pine in a set of morphological and physiological traits: soil-to-leaf hydraulic conductance, intrinsic water-use efficiency (WUE, estimated by carbon isotope composition, δ(13)C), root morphology, xylem anatomy, growth and carbon allocation patterns. The data were collected from Pinus pinaster Aiton seedlings (25 half-sib families from five populations) grown in a greenhouse and subjected to water and water-stress treatments. The aims were to relate this variability to differences in water availability at the geographic location of the populations, and to study the potential trade-offs among traits. The drought-stressed seedlings demonstrated a decrease in hydraulic conductance and root surface area and increased WUE and root tip number. The relationships among the growth, morphological, anatomical and physiological traits changed with the scale of study: within the species, among/within populations. The populations showed a highly significant relationship between the percentage reduction in whole-plant hydraulic conductance and WUE. The differences among the populations in root morphology, whole-plant conductance, carbon allocation, plant growth and WUE were significant and consistent with dryness of the site of seed origin. The xeric populations exhibited lower growth and a conservative water use, as opposed to the fast-growing, less water-use-efficient populations from mesic habitats. The xeric and mesic populations, Tamrabta and San Cipriano, respectively, showed the most contrasting traits and were clustered in opposite directions along the main axis in the canonical discriminant analysis under both the control and drought treatments. The results suggest the possibility of selecting the Arenas population, which presents a combination of traits that confer increased growth and drought resistance.

The present study attempts to investigate the size, composition and seedling dynamics of the seed bank of a metalliferous and vulnerable species, Erica andevalensis. Samples were taken during spring and autumn from two different sites. We also studied the effects of nutrient solution, irrigation from the river Tinto and irrigation deficit on seeding establishment and survival. Only E. andevalensis and Poaceae species emerged from the seed banks, although the former was dominant (98%). Germination and seedling establishment was totally inhibited by the waters of the river Tinto. Seed density was high in the soils of both seed banks irrigated with water and nutrient solution. We found no seasonal differences in the seed bank and number of germinated seeds and mortality rate and density were similar. From the standpoint of restoration management, the results indicate that the seed bank is a very important factor for successful species establishment., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Juniperus thurifera L. is an endemic conifer of the western Mediterranean Basin where it is subjected to a severe climatic stress characterized by low winter temperatures and summer drought. Given the trend of increased warming-induced drought stress in this area and the climatic sensitivity of this species, we expect a negative impact of climate change on growth and ecophysiological performance of J. thurifera in the harsh environments where it dominates. To evaluate this, we measured long- and short-term radial growth using dendrochronology, photosynthesis and water-use efficiency in males, females and juveniles in three sites in Central Spain. Climate was monitored and completed with historical records. Mean annual temperature has increased +0.2 °C per decade in the study area, and the main warming trends corresponded to spring (+0.2 °C per decade) and summer (+0.3 °C per decade). Radial growth and maximum photosynthesis peaked in spring and autumn. Positive photosynthetic rates were maintained all year long, albeit at reduced rates in winter and summer. Radial growth was enhanced by wet conditions in the previous autumn and by warm springs and high precipitation in summer of the year of tree-ring formation. Cloud cover during the summer increased growth, while cloudy winters led to impaired carbon gain and reduced growth in the long term. We argue that maintenance of carbon gain under harsh conditions (low winter temperatures and dry summer months) and plastic xylogenesis underlie J. thurifera's ability to profit from changing climatic conditions such as earlier spring onset and erratic summer rainfall. Our results highlight that not only the magnitude but also the sign of the impact of climate change on growth and persistence of Mediterranean trees is species specific.

Strobilurins are one of the most important classes of agricultural fungicides. In addition to their anti-fungal effect, strobilurins have been reported to produce simultaneous effects in plant physiology. This study investigated whether the use of strobilurin fungicide improved water use efficiency in leaves of grapevines grown under field conditions in a Mediterranean climate in southern Spain. Fungicide was applied three times in the vineyard and measurements of leaf gas exchange, plant water status, abscisic acid concentration in sap ([ABA]), and carbon isotope composition in leaves were performed before and after applications. No clear effect on stomatal conductance, leaf water potential and intrinsic water use efficiency was found after three fungicide applications. ABA concentration was observed to increase after fungicide application on the first day, vanishing three days later. Despite this transient effect, evolution of [ABA] matched well with the evolution of leaf carbon isotope ratio, which can be used as a surrogate for plant water use efficiency. Morning stomatal conductance was negatively correlated to [ABA]. Yield was enhanced in strobilurin treated plants, whereas fruit quality remained unaltered., (Published by Elsevier GmbH.)

Silvicultural thinning usually improves the water status of remaining trees in water-limited forests. We evaluated the usefulness of a dual stable isotope approach (δ¹³C, δ¹⁸O) for comparing the physiological performance of remaining trees between forest stands subjected to two different thinning intensities (moderate versus heavy) in a 60-year-old Pinus halepensis Mill. plantation in semiarid southeastern Spain. We measured bulk leaf δ¹³C and δ¹⁸O, foliar elemental concentrations, stem water content, stem water δ¹⁸O (δ¹⁸O(stem water)), tree ring widths and leaf gas exchange rates to assess the influence of forest stand density on tree performance. Remaining trees in low-density stands (heavily thinned) showed lower leaf δ¹⁸O, and higher stomatal conductance (g(s)), photosynthetic rate and radial growth than those in moderate-density stands (moderately thinned). By contrast, leaf δ¹³C, intrinsic water-use efficiency, foliar elemental concentrations and δ¹⁸O(stem water) were unaffected by stand density. Lower foliar δ¹⁸O in heavily thinned stands reflected higher g(s) of remaining trees due to decreased inter-tree competition for water, whereas higher photosynthetic rate was largely attributable to reduced stomatal limitation to CO₂ uptake. The dual isotope approach provided insight into the early (12 months) effects of stand density manipulation on the physiological performance of remaining trees., (© 2011 Blackwell Publishing Ltd.)

Plant growth response to salinity on a scale of years has not been studied in terms of growth analysis. To gain insights into this topic, 2-year-old Mediterranean Fan Palm (Chamaerops humilis L.) and Mexican Fan Palm (Washingtonia robusta H. Wendl) seedlings, each with its own distinct plant morphology, were grown for 2 years in a peat soil and irrigated with water of 2 dS m(-1) (control) or 8 dS m(-1) (saline). Plants were harvested on seven occasions and the time trends in relative growth rate (RGR, the rate of increase of biomass per unit of biomass already existing) and its components were analysed. In the long term, salinity produced a slight reduction in the mean RGR, values in both species. In the short term, salinity caused a reduction in RGR. However, during the second year, plants irrigated with 8 dS m(-1) grew somewhat more quickly than the control plants, probably as a result of delay in the growth kinetics due to salinity. Regarding RGR components, leaf nitrogen productivity (the rate of biomass gain per unit leaf N and time) was the major factor causing the differences in RGR resulting from salinity. Washingtonia robusta showed a relatively high plasticity in plant morphology by increasing root and decreasing stem biomass allocation in the presence of salinity. However, the long-term response of W. robusta to salinity, based to a great extent, on this morphological plasticity, was less effective than that of C. humilis, which is based mainly on the contribution of leaf N to RGR values.

Little is known about how environmental factors shape the short- and long-term responses of leaf respiration to temperature under field conditions despite the importance of respiration for plant and stand carbon balances. Impacts of water availability and canopy cover on leaf dark respiration (R) and temperature sensitivity were assessed in beech (Fagus sylvatica L.) seedlings in a sub-Mediterranean population. We studied seedlings established within canopy gaps (39% global site factor; GSF) that were subject to either no watering (unwatered plants; UW) or regular watering (2-10% higher volumetric topsoil water content as summer progressed; W plants) and seedlings established beneath the adjacent understorey (12% GSF). Leaf R rose exponentially with diurnal increases in temperature; the same temperature sensitivity (Q(10): 2.2) was found for understorey and gap plants, irrespective of watering treatment. Respiration estimated at 25 degrees C (R(25)) was lower in the understorey than the gaps and was significantly lower in the unwatered than in the watered gap plants by the end of summer (0.65 versus 0.80 micromol m(-2) s(-1)). R(25) declined with increasing summer temperature in all plants; however, respiration estimated at the prevailing ambient temperature did not change through the summer. There were parallel declines in R(25) and concentrations of starch and soluble sugars with increasing summer temperature for gap plants. We conclude that seasonal shifts in temperature-response curves of beech leaf R occur in both low- and high-light environments; since leaf R decreased with increasing plant water deficit, such shifts are likely to be greater whenever plants experience summer drought compared to scenarios where plants experience high rainfall in summer.

Post-anthesis leaf senescence is a key developmental process in the life of plants as it is the time during which material built up by the plant during its growth phase is mobilized into reproductive tissues. Here we aimed to study the extent of photo- and antioxidant protection and salicylic acid (SA) accumulation during post-anthesis leaf senescence in a perennial plant, Salvia lanigera Poir. grown under Mediterranean field conditions. SA levels increased sharply (up to 2.7-fold) during early stages of leaf senescence until fruit and seed formation occurred (i.e. 4 weeks after anthesis). Later on, SA levels kept at constant high levels until leaf abscission occurred (i.e. 7 weeks after anthesis). Reductions in chlorophyll and carotenoid (lutein, violaxanthin and beta-carotene) levels occurred progressively during leaf senescence. In contrast, xanthophyll cycle de-epoxidation increased during early stages of leaf senescence and remained constant later, similar to SA accumulation. Indeed, xanthophyll cycle de-epoxidation strongly positively correlated with SA levels (r(2) = 0.92). The maximum efficiency of PSII (F(v)/F(m) ratio) kept around 0.80 throughout the experiment, except during the latest stage of leaf senescence (i.e. after fruit and seed formation), when this ratio decreased to 0.72, thus indicating damage to PSII. It is concluded that endogenous SA levels increase sharply during early stages of post-anthesis leaf senescence and concomitantly with activation of photoprotection mechanisms, such as xanthophyll cycle-dependent excess energy dissipation, thus avoiding damage to PSII until fruit and seed formation have been accomplished.

We studied photoprotection and antioxidative protection in the three major species of the Canarian laurel forest (Laurus azorica (Seub.) Franco, Persea indica (L.) K. Spreng and Myrica faya Aiton). Trees were exposed to drought under controlled conditions by withholding water until leaf relative water content (RWC) reached 50-55%. Drought reduced photosynthetic rate (P(N)) and was associated with decreased quantum yield of photosystem II (PSII) electron transport and increased non-photochemical quenching (NPQ) in L. azorica and M. faya, but did not increase NPQ in P. indica. Drought-treated trees of L. azorica had the highest de-epoxidation state (DPS) of the xanthophyll cycle and the highest zeaxanthin (Z) concentration, suggesting that this species had more effective photoprotective mechanisms than M. faya and P. indica. Moreover, beta-carotene remained unaltered in L. azorica trees during drought, suggesting that the chloroplasts of this species are better protected against oxidative stress than those of M. faya and P. indica. Increased antioxidation by ascorbate peroxidase, superoxide dismutase and glutathione reductase in L. azorica removed activated oxygen species (AOS) generated during drought treatment. Although M. faya was able to increase its energy dissipation rate by forming Z and thus increasing the DPS of the xanthophyll cycle, it did not respond to drought-induced oxidative stress with the result that beta-carotene degradation occurred. Persea indica did not activate an energy dissipation mechanism in response to drought treatment, hence formation of AOS was likely high in the drought-treated trees. In general, L. azorica was most resistant and P. indica most sensitive to photoinhibition and oxidative stress during drought.

Here, we investigated the physiological and structural leaf responses of seedlings of two evergreen and two deciduous Quercus species, grown in a glasshouse and subjected to contrasted conditions of light (low, medium and high irradiance) and water (continuous watering vs 2-months drought). The impact of drought on photosynthetic rate was strongest in high irradiance, while the impact of shade on photosynthetic rate was strongest with high water supply, contradicting the hypothesis of allocation trade-off. Multivariate causal models were evaluated using d-sep method. The model that best fitted the dataset proposed that the variation in specific leaf area affects photosynthetic rate and leaf nitrogen concentration, and this trait determines stomatal conductance, which also affects photosynthetic rate. Shade conditions seemed to ameliorate, or at least not aggravate, the drought impact on oak seedlings, therefore, the drought response on leaf performance depended on the light environment.

Quercus ilex L. subsp. ballota (Desf.) Samp., a Mediterranean evergreen species growing in a continental Mediterranean climate, did not experience water stress and showed greater sensitivity to winter stress than to summer stress over a 12-month period. Net CO2 assimilation rates and photosystem II (PSII) efficiency decreased markedly during the cold months and recovered completely in spring. Lutein, neoxanthin and beta-carotene to chlorophyll (Chl) molar ratios all showed the same trend throughout the year, increasing from September to March. This increase was a result of increases in carotenoid concentrations, because Chl concentration per unit leaf area remained stable, and was higher at the end than at the beginning of the first growing season. Lutein-epoxide was a minor component of the total lutein pool. Thermal energy dissipation and non-photochemical quenching (NPQ) were associated with the de-epoxidated forms of the xanthophyll cycle pigments in the warm months. Photosynthetic rates decreased slightly at midday in summer. These changes were accompanied by decreases in maximum potential PSII efficiency (which recovered during the night), actual and intrinsic PSII efficiencies, photochemical quenching and increases in NPQ. Overall, our data indicate down-regulation of photosynthesis during the summer. The diurnal de-epoxidation of violaxanthin to antheraxanthin and zeaxanthin occurred throughout the year, except in January. Antioxidant enzymatic activity increased in the winter months, especially during the coldest months, highlighting its key role in photoprotection against photo-oxidation. Structural and functional modifications protected PSII from permanent damage and allowed 1-year-old leaves to photosynthesize at high rates when temperatures increased in spring.

We studied the interspecific and intraspecific variation in the development of water stress and in the use of different water and nitrogen sources during the spring (wet season) and summer (dry season) in a shrub community in NE Spain. We measured shoot water potentials, stable deuterium isotopic composition (delta D) of xylem sap, leaf mass per area, leaf N and C concentrations, gas exchange, leaf delta(13)C, and leaf delta(15)N of the dominant species (Quercus coccifera, Arbutus unedo, Pistacia lentiscus, Erica multiflora, Globularia alypum). The delta D, the delta(13)C and the shoot water potential values showed diurnal, seasonal, intraspecific and interspecific variation in the source and use of water. There was also seasonal, intraspecific and interspecific variation in the foliar delta(15)N and N concentrations. In summer, some species (A. unedo, P. lentiscus and E. multiflora) presented significantly different delta D values in morning and afternoon measurements likely indicating that they used different sources of water during the day, and a dual root system in these species. We conjecture that dew may be one of these water sources. Species predawn water potential was negatively correlated with species xylem water delta D. There was also a positive correlation between delta(13)C and delta D in P. lentiscus, species for which we took additional samples from nearby sites. These results suggest that the access to water from greater depths allowed the maintenance of more favourable plant water supply. Multivariate principal component analysis based on the studied hydrological and isotope variables clearly separated the seasons (wet spring and dry summer) and the species. The species resulted separated according to their evolutionary history (Pre-Mediterranean and Mediterranean) and the associated root and functional traits. These results show water (and nitrogen) partitioning among coexisting species of the same functional type (Mediterranean woody shrubs). They also show the great intraspecific plasticity of responses to resource availability.