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The Mediterranean will be one of the focal points of climate change. The predicted dry and hot summers will lead to water scarcity in agriculture, which may limit crop production and growth. The olive tree serves as a model woody plant for studying drought stress and improving water resource management; thus, it is critical to identify genotypes that are more drought tolerant and perform better under low irrigation or even rainfed conditions. In this study, the metabolomic approach was used to highlight variations in metabolites in stems and leaves of three Italian olive cultivars (previously characterized physiologically) under two and four weeks of drought stress. Phenolic and lipophilic profiles were obtained by gas chromatography-mass spectrometry and ultra-high performance liquid chromatography-mass spectrometry, respectively. The findings identified the leaf as the primary organ in which phenolic variations occurred. The Maurino cultivar exhibited a strong stress response in the form of phenolic compound accumulation, most likely to counteract oxidative stress. The phenolic compound content of ‘Giarraffa’ and ‘Leccino’ plants remained relatively stable whether they were exposed to drought or not. Variations in the lipid profile occurred in leaves and stems of all the cultivars. A high accumulation of compounds related to epicuticular wax components was observed in the leaf of ‘Giarraffa’, while a strong reduction of lipids and long-chain alkanes occurred in ‘Maurino’ when exposed to drought stress conditions. [ABSTRACT FROM AUTHOR]

Water scarcity in agriculture can limit crop production and trigger the need for more effective water resource management. As a result, it is critical to identify new crop genotypes that are more drought tolerant and perform better under low irrigation or even rain-fed conditions. The olive tree is a high-value crop that is well adapted to dry Mediterranean conditions. However, different genotypes may have developed different mechanisms of tolerance to water stress. To investigate such mechanisms, we examined three Italian olive cultivars ('Giarraffa', 'Leccino', and 'Maurino') grown in a greenhouse under drought stress. We found that single genotypes responded differently to the drought, though not all parameters revealed significant differences. The first major difference among the cultivars was in transpiration: the lower stomatal density and stomatal conductance of 'Giarraffa' allow this cultivar to use water more conservatively. In parallel with the reduction in stomatal and mesophyll conductance, the drought-stressed group of 'Giarraffa' maintained the electron transport rate and effective efficiency levels of photosystem II similar to those of the control until the fourth week of stress. The fluorescence parameters revealed the earlier closure of reaction photosynthetic centres in 'Leccino'. Finally, the higher rate of electrolyte leakage in 'Maurino' indicated a significant ions loss in this cultivar when it was subjected to the drought. Both water management under stress conditions and the effect of drought on photosynthesis make 'Giarraffa' interesting to researchers studying its use in breeding or water-saving programmes. [ABSTRACT FROM AUTHOR]

Main conclusion: A water-deficit period, leading to stomatal control and overexpression of protective proteins (sHSP and DHN), contributes to olive´s tolerance to later imposed stress episodes. Aquaporins modulation is important in olive recovery. Olive is traditionally cultivated in dry farming or in high water demanding irrigated orchards. The impact of climate change on these orchards remains to unveil, as heat and drought episodes are increasing in the Mediterranean region. To understand how young plants face such stress episodes, olive plants growing in pots were exposed to well-irrigated and non-irrigated treatments. Subsequently, plants from each treatment were either exposed to 40 °C for 2 h or remained under control temperature. After treatments, all plants were allowed to grow under well-irrigated conditions (recovery). Leaves were compared for photosynthesis, relative water content, mineral status, pigments, carbohydrates, cell membrane permeability, lipid peroxidation and expression of the protective proteins' dehydrin (OeDHN1), heat-shock proteins (OeHSP18.3), and aquaporins (OePIP1.1 and OePIP2.1). Non-irrigation, whilst increasing carbohydrates, reduced some photosynthetic parameters to values below the ones of the well-irrigated plants. However, when both groups of plants were exposed to heat, well-irrigated plants suffered more drastic decreases of net CO2 assimilation rate and chlorophyll b than non-irrigated plants. Overall, OeDHN1 and OeHSP18.3 expression, which was increased in non-irrigated treatment, was potentiated by heat, possibly to counteract the increase of lipid peroxidation and loss of membrane integrity. Plants recovered similarly from both irrigation and temperature treatments, and recovery was associated with increased aquaporin expression in plants exposed to one type of stress (drought or heat). These data represent an important contribution for further understanding how dry-farming olive will cope with drought and heat episodes. [ABSTRACT FROM AUTHOR]

The depletion of the stratospheric ozone layer due to natural and/or anthropogenic causes decreases the amount of UV-B radiation filtered, and consequently increases the risk of potential damage to organisms. In the Mediterranean region, high UV-B indices are frequent. Even for species typical of this region, such as the olive tree, the progressive increase in UV-B radiation represents a threat. This work aimed to understand how high UV-B radiation modulates the phenolic and lipophilic profile of olive varieties, and identify metabolites that enhance olive stress tolerance. Two Italian olive varieties were subjected to chronic UV-B stress, and leaves were analyzed by gas and liquid chromatography. The results indicated that the most representative phenolic and lipophilic compounds of Giarraffa and Olivastra Seggianese were readjusted in response to UV-B stress. The Giarraffa variety seemed better suited to prolonged UV-B stress, possibly due to the higher availability of flavonoids that could help control oxidative damage, and the accumulation of hydroxycinnamic acid derivatives that could provide strong UV-B shield protection. In addition, this variety contained higher levels of fatty acids (e.g., palmitic, α-linolenic, and stearic acids), which can help to maintain membrane integrity and accumulate more sorbitol (which may serve as an osmoprotectant or act a free-radical scavenger), terpenes, and long-chain alkanes, providing higher protection against UV-B stress. [ABSTRACT FROM AUTHOR]

In recent decades, atmospheric pollution led to a progressive reduction of the ozone layer with a consequent increase in UV-B radiation. Despite the high adaptation of olive trees to the Mediterranean environment, the progressive increase of UV-B radiation is a risk factor for olive tree cultivation. It is therefore necessary to understand how high levels of UV-B radiation affect olive plants and to identify olive varieties which are better adapted. In this study we analyzed two Italian olive varieties subjected to chronic UV-B stress. We focused on the effects of UV-B radiation on RubisCO, in terms of quantity, enzymatic activity and isoform composition. In addition, we also analyzed changes in the activity of antioxidant enzymes (SOD, CAT, GPox) to get a comprehensive picture of the antioxidant system. We also evaluated the effects of UV-B on the enzyme sucrose synthase. The overall damage at biochemical level was also assessed by analyzing changes in Hsp70, a protein triggered under stress conditions. The results of this work indicate that the varieties (Giarraffa and Olivastra Seggianese) differ significantly in the use of specific antioxidant defense systems, as well as in the activity and isoform composition of RubisCO. Combined with a different use of sucrose synthase, the overall picture shows that Giarraffa optimized the use of GPox and opted for a targeted choice of RubisCO isoforms, in addition to managing the content of sucrose synthase, thereby saving energy during critical stress points. [ABSTRACT FROM AUTHOR]

The frequency of combined stress events is increasing due to climate change and represents a new threat to olive (Olea europaea) culture. How olive plants modulate their profile of metabolites under multiple stressing agents remains to unveil, although several metabolites affect plants' resilience, and olive production and quality. Young olive plants were exposed to a water deficit (WD) for 30 days and then exposed to a shock of heat and high UVB-radiation (WDHS+UVB treatment) for 2 days. Then, plants were re-watered and grown under optimal conditions (recovery) for 30 days. Leaves were collected after stress and recovery, analysed by liquid and gas chromatography, and the lipophilic and phenolic profiles were characterized. Except for the oleuropein derivatives, the qualitative metabolite profile was similar during stress and recovery. Metabolite increases or decreases in response to stress were stronger when WD was followed by WDHS+UVB treatment. Phenolic compounds (luteolin-7- O -glucoside, quercetin-3- O -rutinoside, apigenin-7- O -glucoside, chrysoeriol-7- O -glucoside, kaempferol derivatives, oleuropein, and lucidumoside C) were the most involved after WD and WDHS+UVB, possibly acting as reactive oxygen species (ROS) scavengers. Lipophilic compounds were more relevant during the recovery period. The catabolism of fatty acids and carbohydrates may provide the necessary energy for plant performance reestablishment, and sterols, long-chain alkanes, and terpenes metabolic pathways may be shifted for the production of compounds with a more important stress protection role. This work highlights for the first time that tolerance mechanisms activated by WD in olive plants are related to metabolite changes, that are adjusted when other stressors are overlapped (WDHS+UVB), and also help the plants recover. This metabolites' plasticity represents an essential contribution to understanding how dry-farming olive orchards may deal with drought combined with high UV-B or heat. [Display omitted] • WD and WDHS+UVB activate the same metabolite responses. • WDHS+UVB induce a stronger metabolite quantity change. • Polyphenols are the most involved to counteract the stress negative effects. • Lipophilic metabolites are more involved in physiological reestablishment after stress. [ABSTRACT FROM AUTHOR]

Extreme climate events are increasingly frequent, and the 2017 summer was particularly critical in the Mediterranean region. Olive is one of the most important species of this region, and these climatic events represent a threat to this culture. However, it remains unclear how olive trees adjust the antioxidant enzymatic system and modulate the metabolite profile under field stress conditions. Leaves from two distinct adjacent areas of an olive orchard, one dry and the other hydrated, were harvested. Tree water status, oxidative stress, antioxidant enzymes, and phenolic and lipophilic metabolite profiles were analyzed. The environmental conditions of the 2017 summer caused a water deficit in olive trees of the dry area, and this low leaf water availability was correlated with the reduction of long-chain alkanes and fatty acids. Hydrogen peroxide (H2O2) and superoxide radical (O2•–) levels increased in the trees collected from the dry area, but lipid peroxidation did not augment. The antioxidant response was predominantly marked by guaiacol peroxidase (GPOX) activity that regulates the H2O2 harmful effect and by the action of flavonoids (luteolin-7-O-glucuronide) that may act as reactive oxygen species scavengers. Secoiridoids adjustments may also contribute to stress regulation. This work highlights for the first time the protective role of some metabolite in olive trees under field drought conditions. [ABSTRACT FROM AUTHOR]

Given the economical importance of the olive tree it is essential to study its responses to stress agents such as excessive UV-B radiation, to understand the defense mechanisms and to identify the varieties that are able to cope with it. In the light of the analysis carried out in this study, we argue that UV-B radiation represents a dangerous source of stress for the olive tree, especially in the current increasingly changing environmental conditions. Both the varieties considered (Giarraffa and Olivastra Seggianese), although resistant to the strong treatment to which they were exposed, showed, albeit in different ways and at different times, evident effects. The two varieties have different response times and the Giarraffa variety seems better suited to prolonged UV-B stress, possible due to a more efficient and quick activation of the antioxidant response (e.g., flavonoids use to counteract reactive oxygen species) and because of its capacity to maintain the photosynthetic efficiency as well as a relatively higher content of mannitol. Moreover, pigments reduction after a long period of UV-B exposure can also be an adaptation mechanism triggered by Giarraffa to reduce energy absorption under UV-B stress. Olivastra Seggianese seems less suited to overcome UV-B stress for a long period (e.g., higher reduction of Fv/Fm) and has a higher requirement for sugars (e.g., glucose) possible to counteract stress and to restore energy. [ABSTRACT FROM AUTHOR]

The Mediterranean crop Olea europaea is often exposed to high UV-B irradiation conditions. To understand how this species modulates its enzymatic and non-enzymatic antioxidant system under high UV-B radiation, young O. europaea plants (cultivar "Galega Vulgar") were exposed, for five days, to UV-B radiation (6.5 kJ m−2 d−1 and 12.4 kJ m−2 d−1). Our data indicate that UV-doses slightly differ in the modulation of the antioxidant protective mechanisms. Particularly, superoxide dismutase (SOD), guaiacol peroxidase (GPox) and catalase (CAT) activities increased contributing to H 2 O 2 homeostasis, being more solicited by higher UV-B doses. Also, glutathione reductase (Gr) activity, ascorbate (AsA) and reduced glutathione (GSH) pools increased particularly under the highest dose, suggesting a higher mobilization of the antioxidant system in this dose. The leaf metabolites' profile of this cultivar was analysed by UHPLC-MS. Interestingly, high levels of verbascoside were found, followed by oleuropein and luteolin-7- O -glucoside. Both UV-B treatments affected mostly less abundant flavonoids (decreasing 4ʹ-methoxy luteolin and 4ʹ or 3ʹ-methoxy luteolin glucoside) and hydroxycinnamic acid derivatives (HCAds, increasing β-hydroxyverbascoside). These changes show not only different mobilization with the UV-intensity, but also reinforce for the first time the protective roles of these minor compounds against UV-B, as reactive oxygen species (ROS) scavengers and UV-B shields, in complement with other antioxidant systems (e.g. AsA/GSH cycle), particularly for high UV-B doses. Secoiridoids also standout in the response to both UV-B doses, with decreases of oleuropein and increases 2ʹʹ-methoxyoleuropein. Being oleuropein an abundant compound, data suggest that secoiridoids play a more important role than flavonoids and HCAds, in O. europaea protection against UV-B, possibly by acting as signalling molecules and ROS scavengers. This is the first report on the influence of UV-B radiation on the secoiridoid oleuropein, and provides a novel insight to the role of this compound in the O. europaea antioxidant defence mechanisms. Image 1 • Moderate and high UV-B doses activated similar antioxidant protective mechanisms. • High UV-B dose promote higher activation of the antioxidant enzymes and AsA/GSH. • Flavonoids, secoiridoids and HCAds respond similarly to 6.5 and 12.4 kJ m−2d−1 UV-B • UV-B treatments reduce some protective compounds level, such as oleuropein. [ABSTRACT FROM AUTHOR]

• Cobrançosa olives are richer in organic acids, esters and carbohydrates. • C.Serpa olives present higher levels of phenolic compounds. • Stress modulates olives' phenolic/lipophilic contents in a cultivar-dependent way. • C.Serpa olives have higher stress susceptibility than Cobrançosa and C.C.Branco. • Olives can be enriched in bioactive compounds in response to environmental stresses. The awareness of the functional and nutraceutical properties of olives and olive oil bioactive constituents contributed to oliviculture recent increase. Olives' metabolism and nutritional quality are determined by how the olive-tree is coping to climate change-related episodes, which increasingly occur in the Mediterranean. We characterize the most relevant lipophilic and phenolic compounds of olives from Olea europaea cultivars [Cobrançosa, Cordovil de Castelo Branco and Cordovil de Serpa (C.Serpa)] exposed to drought + heat. Olives from the three cultivars presented a similar qualitative profile but differed in their relative richness. Cobrançosa olives are richer in organic acids, esters and carbohydrates, while C.Serpa olives have higher levels of phenolic compounds, particularly under control conditions. Drought + heat changed the quantitative profile of olives, in a way dependent on the cultivar, and C.Serpa olives showed the highest stress susceptibility. Climate change-related conditions stimulate the accumulation of relevant bioactive compounds in olives, contributing to increasing its nutritional value. [ABSTRACT FROM AUTHOR]